Improving Auditory Filter Estimation by Incorporating Absolute Threshold and a Level-dependent Internal Noise.

Auditory filter (AF) shape has traditionally been estimated with a combination of a notched-noise (NN) masking experiment and a power spectrum model (PSM) of masking. However, there are several challenges that remain in both the simultaneous and forward masking paradigms. We hypothesized that AF shape estimation would be improved if absolute threshold (AT) and a level-dependent internal noise were explicitly represented in the PSM. To document the interaction between NN threshold and AT in normal hearing (NH) listeners, a large set of NN thresholds was measured at four center frequencies (500, 1000, 2000, and 4000 Hz) with the emphasis on low-level maskers. The proposed PSM, consisting of the compressive gammachirp (cGC) filter and three nonfilter parameters, allowed AF estimation over a wide range of frequencies and levels with fewer coefficients and less error than previous models. The results also provided new insights into the nonfilter parameters. The detector signal-to-noise ratio (K) was found to be constant across signal frequencies, suggesting that no frequency dependence hypothesis is required in the postfiltering process. The ANSI standard "Hearing Level-0dB" function, i.e., AT of NH listeners, could be applied to the frequency distribution of the noise floor for the best AF estimation. The introduction of a level-dependent internal noise could mitigate the nonlinear effects that occur in the simultaneous NN masking paradigm. The new PSM improves the applicability of the model, particularly when the sound pressure level of the NN threshold is close to AT.

Early cortical processing of pitch height and the role of adaptation and musicality.

Pitch is an important perceptual feature; however, it is poorly understood how its cortical correlates are shaped by absolute vs relative fundamental frequency (f0), and by neural adaptation. In this study, we assessed transient and sustained auditory evoked fields (AEFs) at the onset, progression, and offset of short pitch height sequences, taking into account the listener's musicality. We show that neuromagnetic activity reflects absolute f0 at pitch onset and offset, and relative f0 at transitions within pitch sequences; further, sequences with fixed f0 lead to larger response suppression than sequences with variable f0 contour, and to enhanced offset activity. Musical listeners exhibit stronger f0-related AEFs and larger differences between their responses to fixed vs variable sequences, both within sequences and at pitch offset. The results resemble prominent psychoacoustic phenomena in the perception of pitch contours; moreover, they suggest a strong influence of adaptive mechanisms on cortical pitch processing which, in turn, might be modulated by a listener's musical expertise.

Transient and sustained processing of musical consonance in auditory cortex and the effect of musicality.

In recent years, electroencephalography and magnetoencephalography (MEG) have both been used to investigate the response in human auditory cortex to musical sounds that are perceived as consonant or dissonant. These studies have typically focused on the transient components of the physiological activity at sound onset, specifically, the N1 wave of the auditory evoked potential and the auditory evoked field, respectively. Unfortunately, the morphology of the N1 wave is confounded by the prominent neural response to energy onset at stimulus onset. It is also the case that the perception of pitch is not limited to sound onset; the perception lasts as long as the note producing it. This suggests that consonance studies should also consider the sustained activity that appears after the transient components die away. The current MEG study shows how energy-balanced sounds can focus the response waves on the consonance-dissonance distinction rather than energy changes and how source modeling techniques can be used to measure the sustained field associated with extended consonant and dissonant sounds. The study shows that musical dyads evoke distinct transient and sustained neuromagnetic responses in auditory cortex. The form of the response depends on both whether the dyads are consonant or dissonant and whether the listeners are musical or nonmusical. The results also show that auditory cortex requires more time for the early transient processing of dissonant dyads than it does for consonant dyads and that the continuous representation of temporal regularity in auditory cortex might be modulated by processes beyond auditory cortex.NEW & NOTEWORTHY We report a magnetoencephalography (MEG) study on transient and sustained cortical consonance processing. Stimuli were long-duration, energy-balanced, musical dyads that were either consonant or dissonant. Spatiotemporal source analysis revealed specific transient and sustained neuromagnetic activity in response to the dyads; in particular, the morphology of the responses was shaped by the dyad's consonance and the listener's musicality. Our results also suggest that the sustained representation of stimulus regularity might be modulated by processes beyond auditory cortex.

Direct electrophysiological mapping of human pitch-related processing in auditory cortex.

This work sought correlates of pitch perception, defined by neural activity above the lower limit of pitch (LLP), in auditory cortical neural ensembles, and examined their topographical distribution. Local field potentials (LFPs) were recorded in eight patients undergoing invasive recordings for pharmaco-resistant epilepsy. Stimuli consisted of bursts of broadband noise followed by regular interval noise (RIN). RIN was presented at rates below and above the LLP to distinguish responses related to the regularity of the stimulus and the presence of pitch itself. LFPs were recorded from human cortical homologues of auditory core, belt, and parabelt regions using multicontact depth electrodes implanted in Heschl's gyrus (HG) and Planum Temporale (PT), and subdural grid electrodes implanted over lateral superior temporal gyrus (STG). Evoked responses corresponding to the temporal regularity of the stimulus were assessed using autocorrelation of the evoked responses, and occurred for stimuli below and above the LLP. Induced responses throughout the high gamma range (60-200 Hz) were present for pitch values above the LLP, with onset latencies of approximately 70 ms. Mapping of the induced responses onto a common brain space demonstrated variability in the topographical distribution of high gamma responses across subjects. Induced responses were present throughout the length of HG and on PT, which is consistent with previous functional neuroimaging studies. Moreover, in each subject, a region within lateral STG showed robust induced responses at pitch-evoking stimulus rates. This work suggests a distributed representation of pitch processing in neural ensembles in human homologues of core and non-core auditory cortex.

The distribution and nature of responses to broadband sounds associated with pitch in the macaque auditory cortex.

The organisation of pitch-perception mechanisms in the primate cortex is controversial, in that divergent results have been obtained, ranging from a single circumscribed 'pitch centre' to systems widely distributed across auditory cortex. Possible reasons for such discrepancies include different species, recording techniques, pitch stimuli, sampling of auditory fields, and the neural metrics recorded. In the present study, we sought to bridge some of these divisions by examining activity related to pitch in both neurons and neuronal ensembles within the auditory cortex of the rhesus macaque, a primate species with similar pitch perception and auditory cortical organisation to humans. We demonstrate similar responses, in primary and non-primary auditory cortex, to two different types of broadband pitch above the macaque lower limit in both neurons and local field potential (LFP) gamma oscillations. The majority of broadband pitch responses in neurons and LFP sites did not show equivalent tuning for sine tones.

Neuromagnetic correlates of voice pitch, vowel type, and speaker size in auditory cortex.

Vowel recognition is largely immune to differences in speaker size despite the waveform differences associated with variation in speaker size. This has led to the suggestion that voice pitch and mean formant frequency (MFF) are extracted early in the hierarchy of hearing/speech processing and used to normalize the internal representation of vowel sounds. This paper presents a magnetoencephalographic (MEG) experiment designed to locate and compare neuromagnetic activity associated with voice pitch, MFF and vowel type in human auditory cortex. Sequences of six sustained vowels were used to contrast changes in the three components of vowel perception, and MEG responses to the changes were recorded from 25 participants. A staged procedure was employed to fit the MEG data with a source model having one bilateral pair of dipoles for each component of vowel perception. This dipole model showed that the activity associated with the three perceptual changes was functionally separable; the pitch source was located in Heschl's gyrus (bilaterally), while the vowel-type and formant-frequency sources were located (bilaterally) just behind Heschl's gyrus in planum temporale. The results confirm that vowel normalization begins in auditory cortex at an early point in the hierarchy of speech processing.

The Effect of Peripheral Compression on Syllable Perception Measured with a Hearing Impairment Simulator.

Hearing impaired (HI) people often have difficulty understanding speech in multi-speaker or noisy environments. With HI listeners, however, it is often difficult to specify which stage, or stages, of auditory processing are responsible for the deficit. There might also be cognitive problems associated with age. In this paper, a HI simulator, based on the dynamic, compressive gammachirp (dcGC) filterbank, was used to measure the effect of a loss of compression on syllable recognition. The HI simulator can counteract the cochlear compression in normal hearing (NH) listeners and, thereby, isolate the deficit associated with a loss of compression in speech perception. Listeners were required to identify the second syllable in a three-syllable "nonsense word", and between trials, the relative level of the second syllable was varied, or the level of the entire sequence was varied. The difference between the Speech Reception Threshold (SRT) in these two conditions reveals the effect of compression on speech perception. The HI simulator adjusted a NH listener's compression to that of the "average 80-year old" with either normal compression or complete loss of compression. A reference condition was included where the HI simulator applied a simple 30-dB reduction in stimulus level. The results show that the loss of compression has its largest effect on recognition when the second syllable is attenuated relative to the first and third syllables. This is probably because the internal level of the second syllable is attenuated proportionately more when there is a loss of compression.

Locating Melody Processing Activity in Auditory Cortex with Magnetoencephalography.

This paper describes a technique for isolating the brain activity associated with melodic pitch processing. The magnetoencephalograhic (MEG) response to a four note, diatonic melody built of French horn notes, is contrasted with the response to a control sequence containing four identical, "tonic" notes. The transient response (TR) to the first note of each bar is dominated by energy-onset activity; the melody processing is observed by contrasting the TRs to the remaining melodic and tonic notes of the bar (2-4). They have uniform shape within a tonic or melodic sequence which makes it possible to fit a 4-dipole model and show that there are two sources in each hemisphere--a melody source in the anterior part of Heschl's gyrus (HG) and an onset source about 10 mm posterior to it, in planum temporale (PT). The N1m to the initial note has a short latency and the same magnitude for the tonic and the melodic sequences. The melody activity is distinguished by the relative sizes of the N1m and P2m components of the TRs to notes 2-4. In the anterior source a given note elicits a much larger N1m-P2m complex with a shorter latency when it is part of a melodic sequence. This study shows how to isolate the N1m, energy-onset response in PT, and produce a clean melody response in the anterior part of auditory cortex (HG).

Disruption of the auditory response to a regular click train by a single, extra click.

It has been hypothesized that the steady-state response to a periodic sequence of clicks can be modeled as the superposition of responses to single clicks. Here, this hypothesis is challenged by presenting an extra click halfway between two consecutive clicks of a regular series, while measuring the auditory evoked field. After a solitary click at time zero, the click series sounded from 100 to 900 ms, with the extra click presented around 500 ms. The silent period between two stimulus sequences was 310-390 ms (uniformly distributed) so that one stimulation cycle lasted, on average, 1250 ms. Five different click rates between 20 and 60 Hz were examined. The disturbance caused by the extra click was revealed by subtracting the estimated steady-state response from the joint response to the click series and the extra click. The early peaks of the single-click response effectively coincide with same-polarity peaks of the 20-Hz steady-state response. Nevertheless, prediction of the latter from the former proved impossible. However, the 40-Hz steady-state response can be predicted reasonably well from the 20-Hz steady-state response. Somewhat surprisingly, the amplitude of the evoked response to the extra click grew when the click rate of the train was increased from 20 to 30 Hz; the opposite effect would have been expected from research on adaptation. The smaller amplitude at lower click rates might be explained by forward suppression. In this case, the apparent escape from suppression at higher rates might indicate that the clicks belonging to the periodic train are being integrated into an auditory stream, possibly in much the same manner as in classical stream segregation experiments.

Tracking cortical entrainment in neural activity: auditory processes in human temporal cortex.

A primary objective for cognitive neuroscience is to identify how features of the sensory environment are encoded in neural activity. Current auditory models of loudness perception can be used to make detailed predictions about the neural activity of the cortex as an individual listens to speech. We used two such models (loudness-sones and loudness-phons), varying in their psychophysiological realism, to predict the instantaneous loudness contours produced by 480 isolated words. These two sets of 480 contours were used to search for electrophysiological evidence of loudness processing in whole-brain recordings of electro- and magneto-encephalographic (EMEG) activity, recorded while subjects listened to the words. The technique identified a bilateral sequence of loudness processes, predicted by the more realistic loudness-sones model, that begin in auditory cortex at ~80 ms and subsequently reappear, tracking progressively down the superior temporal sulcus (STS) at lags from 230 to 330 ms. The technique was then extended to search for regions sensitive to the fundamental frequency (F0) of the voiced parts of the speech. It identified a bilateral F0 process in auditory cortex at a lag of ~90 ms, which was not followed by activity in STS. The results suggest that loudness information is being used to guide the analysis of the speech stream as it proceeds beyond auditory cortex down STS toward the temporal pole.

A perceptual pitch boundary in a non-human primate.

Pitch is an auditory percept critical to the perception of music and speech, and for these harmonic sounds, pitch is closely related to the repetition rate of the acoustic wave. This paper reports a test of the assumption that non-human primates and especially rhesus monkeys perceive the pitch of these harmonic sounds much as humans do. A new procedure was developed to train macaques to discriminate the pitch of harmonic sounds and thereby demonstrate that the lower limit for pitch perception in macaques is close to 30 Hz, as it is in humans. Moreover, when the phases of successive harmonics are alternated to cause a pseudo-doubling of the repetition rate, the lower pitch boundary in macaques decreases substantially, as it does in humans. The results suggest that both species use neural firing times to discriminate pitch, at least for sounds with relatively low repetition rates.

Duifhuis pitch: neuromagnetic representation and auditory modeling.

When a high harmonic is removed from a cosine-phase harmonic complex, we hear a sine tone pop out of the perception; the sine tone has the pitch of the high harmonic, while the tone complex has the pitch of its fundamental frequency, f0. This phenomenon is commonly referred to as Duifhuis Pitch (DP). This paper describes, for the first time, the cortical representation of DP observed with magnetoencephalography. In experiment 1, conditions that produce the perception of a DP were observed to elicit a classic onset response in auditory cortex (P1m, N1m, P2m), and an increment in the sustained field (SF) established in response to the tone complex. Experiment 2 examined the effect of the phase spectrum of the complex tone on the DP activity: Schroeder-phase negative waves elicited a transient DP complex with a similar shape to that observed with cosine-phase waves but with much longer latencies. Following the transient DP activity, the responses of the negative and positive Schroeder-phase waves converged, and the increment in the SF slowly died away. In the absence of DP, the two Schroeder-phase conditions with low peak factors both produced larger SFs than cosine-phase waves with large peak factors. A model of the auditory periphery that includes coupling between adjacent frequency channels is used to explain the early neuromagnetic activity observed in auditory cortex.

Accurate estimation of compression in simultaneous masking enables the simulation of hearing impairment for normal-hearing listeners.

This chapter presents a unified gammachirp framework for -estimating cochlear compression and synthesizing sounds with inverse compression that -cancels the compression of a normal-hearing (NH) listener to simulate the -experience of a hearing-impaired (HI) listener. The compressive gammachirp (cGC) filter was -fitted to notched-noise masking data to derive level-dependent -filter shapes and the cochlear compression function (e.g., Patterson et al., J Acoust Soc Am 114:1529-1542, 2003). The procedure is based on the analysis/synthesis technique of Irino and Patterson (IEEE Trans Audio Speech Lang Process 14:2222-2232, 2006) using a dynamic cGC filterbank (dcGC-FB). The level dependency of the dcGC-FB can be reversed to produce inverse compression and resynthesize sounds in a form that cancels the compression applied by the -auditory system of the NH listener. The chapter shows that the estimation of compression in simultaneous masking is improved if the notched-noise procedure for the derivation of auditory filter shape includes noise bands with different levels. Since both the estimation and resynthesis are performed within the gammachirp framework, it is possible for a specific NH listener to experience the loss of a -specific HI listener.

Modelling the distortion produced by cochlear compression.

Lyon (J Acoust Soc Am 130:3893-3904, 2011) has described how a cascade of simple asymmetric resonators (CAR) can be used to simulate the filtering of the basilar membrane and how the gain of the resonators can be manipulated by a feedback network to simulate the fast-acting compression (FAC) characteristic of cochlear processing. When the compression is applied to complex tones, each pair of primary components produces both quadratic and cubic distortion tones (DTs), and the cascade architecture of the CAR-FAC system propagates them down to their appropriate place along the basilar membrane, where they combine additively with each other and any primary components at that frequency. This suggests that CAR-FAC systems might be used to study the role of compressive distortion in the perception of complex sounds and that behavioural measurements of cochlear distortion data might be useful when tuning the parameters of CAR-FAC systems.

Cortical activity associated with the perception of temporal asymmetry in ramped and damped noises.

Human listeners are very sensitive to the asymmetry of time-reversed pairs of ramped and damped sounds. When the carrier is noise, the hiss -component of the perception is stronger in ramped sounds and the drumming component is stronger in damped sounds (Akeroyd and Patterson 1995). In the current study, a paired comparison technique was used to establish the relative "hissiness" of these noises, and the ratings were correlated with (a) components of the auditory evoked field (AEF) produced by these noises and (b) the magnitude of a hissiness feature derived from a model of the internal auditory images produced by these noises (Irino and Patterson 1998). An earlier AEF report indicated that the peak magnitude of the transient N100m response mirrors the perceived salience of the tonal perception (Rupp et al. 2005). The AEFs of 14 subjects were recorded in response to damped/ramped noises with half-lives between 1 and 64 ms and repetition rates between 12.5 and 100 ms. Spatio-temporal source analysis was used to fit the P50m, the P200m, and the sustained field (SF). These noise stimuli did not produce a reliable N100m. The hissiness feature from the auditory model was extracted from a time-averaged sequence of summary auditory images as in Patterson and Irino (1998). The results show that the perceptual measure of hissiness is highly correlated with the hissiness feature from the summary auditory image, and both are highly correlated with the magnitude of the transient P200m. There is a significant but weaker correlation with the SF and a nonsignificant correlation with the P50m. The results suggest that regularity in the carrier effects branching at an early stage of auditory processing with tonal and noisy sounds following separate spatio-temporal routes through the system.

Age-related difference in melodic pitch perception is probably mediated by temporal processing: empirical and computational evidence.

OBJECTIVE: This study was designed to examine whether age-related differences in melodic pitch perception may be mediated by temporal processing. Temporal models of pitch suggest that performance will decline as the lowest component of a complex tone increases in frequency, regardless of age. In addition, if there are age-related deficits in temporal processing in older adults, this group may have reduced performance relative to younger adults even in the most favorable conditions. DESIGN: Six younger adults and 10 older adults with clinically normal audiograms up to 8 kHz were tested in a melodic pitch perception task. In each trial, two consecutive four-note melodies were presented to the listener. Melodies were identical with the exception of one note in the second melody that was shifted in pitch. The listener was required to identify which note was shifted. All notes consisted of eight successive harmonic components, with the average lowest component manipulated to be the 4th, 8th, or 12th component of the harmonic series, with lower components being absent. RESULTS: Age-related differences in melodic pitch perception were only apparent when stimulus parameters favored temporal processing of pitch. Furthermore, modeling a loss of periodicity coding yielded an outcome consistent with the observed behavioral results. Although younger adults generally outperformed older adults, about one-quarter of the older adults performed at levels that were equivalent to those of younger adults. The only follow-up tests that were able to differentiate these exceptional older adults were tests that would be sensitive to temporal processing: fundamental frequency difference limens and 500 Hz pure-tone difference limens. In contrast, otoacoustic emissions and high-frequency pure-tone thresholds, which are more commonly associated with spectral processing deficits, were not able to differentiate older exceptional adults from older typical adults. CONCLUSION: Age-related declines in temporal processing contribute to deficits in melodic pitch perception. However, some exceptional older adults with normal audiograms preserve excellent temporal processing and continue to perform at levels that are typical of younger adults.

The mutual roles of temporal glimpsing and vocal characteristics in cocktail-party listening.

At a cocktail party, listeners must attend selectively to a target speaker and segregate their speech from distracting speech sounds uttered by other speakers. To solve this task, listeners can draw on a variety of vocal, spatial, and temporal cues. Recently, Vestergaard et al. [J. Acoust. Soc. Am. 125, 1114-1124 (2009)] developed a concurrent-syllable task to control temporal glimpsing within segments of concurrent speech, and this allowed them to measure the interaction of glottal pulse rate and vocal tract length and reveal how the auditory system integrates information from independent acoustic modalities to enhance recognition. The current paper shows how the interaction of these acoustic cues evolves as the temporal overlap of syllables is varied. Temporal glimpses as short as 25 ms are observed to improve syllable recognition substantially when the target and distracter have similar vocal characteristics, but not when they are dissimilar. The effect of temporal glimpsing on recognition performance is strongly affected by the form of the syllable (consonant-vowel versus vowel-consonant), but it is independent of other phonetic features such as place and manner of articulation.

Neuromagnetic representation of musical register information in human auditory cortex.

Pulse-resonance sounds like vowels or instrumental tones contain acoustic information about the physical size of the sound source (pulse rate) and body resonators (resonance scale). Previous research has revealed correlates of these variables in humans using functional neuroimaging. Here, we report two experiments that use magnetoencephalography to study the neuromagnetic representations of pulse rate and resonance scale in human auditory cortex. In Experiment 1, auditory evoked fields were recorded from nineteen subjects presented with French horn tones, the pulse rate and resonance scale of which had been manipulated independently using a mucoder. In Experiment 2, fifteen subjects listened to French horn tones which differed in resonance scale but which lacked pulse rate cues. The resulting cortical activity was evaluated by spatio-temporal source analysis. Changes in pulse rate elicited a well-defined N1m component with cortical generators located at the border between Heschl's gyrus and planum temporale. Changes in resonance scale elicited a second, independent, N1m component located in planum temporale. Our results demonstrate that resonance scale can be distinguished in its neuromagnetic representation from cortical activity related to the sound's pulse rate. Moreover, the existence of two separate components in the N1m sensitive to register information highlights the importance of this time window for the processing of frequency information in human auditory cortex.

Predictive coding and pitch processing in the auditory cortex.

In this work, we show that electrophysiological responses during pitch perception are best explained by distributed activity in a hierarchy of cortical sources and, crucially, that the effective connectivity between these sources is modulated with pitch strength. Local field potentials were recorded in two subjects from primary auditory cortex and adjacent auditory cortical areas along the axis of Heschl's gyrus (HG) while they listened to stimuli of varying pitch strength. Dynamic causal modeling was used to compare system architectures that might explain the recorded activity. The data show that representation of pitch requires an interaction between nonprimary and primary auditory cortex along HG that is consistent with the principle of predictive coding.

Auditory cortex tracks the temporal regularity of sustained noisy sounds.

Neuroimaging studies have revealed dramatic asymmetries between the responses to temporally regular and irregular sounds in the antero-lateral part of Heschl's gyrus. For example, the magnetoencephalography (MEG) study of Krumbholz et al. [Cereb. Cortex 13, 765-772 (2003)] showed that the transition from a noise to a similar noise with sufficient temporal regularity to provoke a pitch evoked a pronounced temporal-regularity onset response (TRon response), whereas a comparable transition in the reverse direction revealed essentially no temporal-regularity offset response (TRoff response). The current paper presents a follow-up study in which the asymmetry is examined with much greater power, and the results suggest an intriguing reinterpretation of the onset/offset asymmetry. The TR-related activity in auditory cortex appears to be composed of a transient (TRon) and a TR-related sustained response (TRsus), with a highly variable TRon/TRsus amplitude ratio. The TRoff response is generally dominated by the break-down of the TRsus activity, which occurs so rapidly as to preclude the involvement of higher-level cortical processing. The time course of the TR-related activity suggests that TR processing might be involved in monitoring the environment and alerting the brain to the onset and offset of behaviourally relevant, animate sources.