Auditory Perception: Laurel and Yanny Together at Last.

An auditory illusion caught the world's attention recently. For the same noisy speech utterance, different people reported hearing either 'Laurel' or 'Yanny'. The dichotomy highlights how perceptions are inferences from inherently ambiguous sensory information, even though ambiguity is often unnoticed.

Interindividual variability in auditory scene analysis revealed by confidence judgements.

Because musicians are trained to discern sounds within complex acoustic scenes, such as an orchestra playing, it has been hypothesized that musicianship improves general auditory scene analysis abilities. Here, we compared musicians and non-musicians in a behavioural paradigm using ambiguous stimuli, combining performance, reaction times and confidence measures. We used 'Shepard tones', for which listeners may report either an upward or a downward pitch shift for the same ambiguous tone pair. Musicians and non-musicians performed similarly on the pitch-shift direction task. In particular, both groups were at chance for the ambiguous case. However, groups differed in their reaction times and judgements of confidence. Musicians responded to the ambiguous case with long reaction times and low confidence, whereas non-musicians responded with fast reaction times and maximal confidence. In a subsequent experiment, non-musicians displayed reduced confidence for the ambiguous case when pure-tone components of the Shepard complex were made easier to discern. The results suggest an effect of musical training on scene analysis: we speculate that musicians were more likely to discern components within complex auditory scenes, perhaps because of enhanced attentional resolution, and thus discovered the ambiguity. For untrained listeners, stimulus ambiguity was not available to perceptual awareness.This article is part of the themed issue 'Auditory and visual scene analysis'.

A Neural Substrate for Rapid Timbre Recognition? Neural and Behavioral Discrimination of Very Brief Acoustic Vowels.

The timbre of a sound plays an important role in our ability to discriminate between behaviorally relevant auditory categories, such as different vowels in speech. Here, we investigated, in the primary auditory cortex (A1) of anesthetized guinea pigs, the neural representation of vowels with impoverished timbre cues. Five different vowels were presented with durations ranging from 2 to 128 ms. A psychophysical experiment involving human listeners showed that identification performance was near ceiling for the longer durations and degraded close to chance level for the shortest durations. This was likely due to spectral splatter, which reduced the contrast between the spectral profiles of the vowels at short durations. Effects of vowel duration on cortical responses were well predicted by the linear frequency responses of A1 neurons. Using mutual information, we found that auditory cortical neurons in the guinea pig could be used to reliably identify several vowels for all durations. Information carried by each cortical site was low on average, but the population code was accurate even for durations where human behavioral performance was poor. These results suggest that a place population code is available at the level of A1 to encode spectral profile cues for even very short sounds.

Adaptive coding is constrained to midline locations in a spatial listening task.

Many neurons adapt their spike output to accommodate the prevailing sensory environment. Although such adaptation is thought to improve coding of relevant stimulus features, the relationship between adaptation at the neural and behavioral levels remains to be established. Here we describe improved discrimination performance for an auditory spatial cue (interaural time differences, ITDs) following adaptation to stimulus statistics. Physiological recordings in the midbrain of anesthetized guinea pigs and measurement of discrimination performance in humans both demonstrate improved coding of the most prevalent ITDs in a distribution, but with highest accuracy maintained for ITDs corresponding to frontal locations, suggesting the existence of a fovea for auditory space. A biologically plausible model accounting for the physiological data suggests that neural tuning is stabilized by inhibition to maintain high discriminability for frontal locations. The data support the notion that adaptive coding in the midbrain is a key element of behaviorally efficient sound localization in dynamic acoustic environments.

Physiological correlates of comodulation masking release in the mammalian ventral cochlear nucleus.

Comodulation masking release (CMR) enhances the detection of signals embedded in wideband, amplitude-modulated maskers. At least part of the CMR is attributable to across-frequency processing, however, the relative contribution of different stages in the auditory system to across-frequency processing is unknown. We have measured the responses of single units from one of the earliest stages in the ascending auditory pathway, the ventral cochlear nucleus, where across frequency processing may take place. A sinusoidally amplitude-modulated tone at the best frequency of each unit was used as a masker. A pure tone signal was added in the dips of the masker modulation (reference condition). Flanking components (FCs) were then added at frequencies remote from the unit best frequency. The FCs were pure tones amplitude modulated either in phase (comodulated) or out of phase (codeviant) with the on-frequency component. Psychophysically, this CMR paradigm reduces within-channel cues while producing an advantage of approximately 10 dB for the comodulated condition in comparison with the reference condition. Some of the recorded units showed responses consistent with perceptual CMR. The addition of the comodulated FCs produced a strong reduction in the response to the masker modulation, making the signal more salient in the poststimulus time histograms. A decision statistic based on d' showed that threshold was reached at lower signal levels for the comodulated condition than for reference or codeviant conditions. The neurons that exhibited such a behavior were mainly transient chopper or primary-like units. The results obtained from a subpopulation of transient chopper units are consistent with a possible circuit in the cochlear nucleus consisting of a wideband inhibitor contacting a narrowband cell. A computational model was used to confirm the feasibility of such a circuit.

The responses of single units in the inferior colliculus of the guinea pig to damped and ramped sinusoids.

Temporal asymmetry can have a pronounced effect on the perception of a sinusoid. For instance, if a sinusoid is amplitude modulated by a decaying exponential that restarts every 50 ms, (a damped sinusoid) listeners report a two-component percept: a tonal component corresponding to the carrier and a drumming component corresponding to the envelope modulation period. When the amplitude modulation is reversed in time (a ramped sinusoid) the perception changes markedly; the tonal component increases while the drumming component decreases. The long-term Fourier energy spectra are identical for damped and ramped sinusoids with the same exponential half-life. Modelling studies suggest that this perceptual asymmetry must occur central to the peripheral stages of auditory processing (Patterson and Irino, 1998). To test this hypothesis, we have recorded the responses of single units in the inferior colliculus of the anaesthetised guinea pig. We divided single units into three groups: onset, on-sustained and sustained, based on their temporal adaptation properties to suprathreshold tone bursts at the unit's best frequency. The asymmetry observed in the neural responses of single units was quantified in two ways: a simple total spike count measure and a ratio of the tallest bin of the modulation period histogram to the total number of spikes. Responses were more diverse than those observed with similar stimuli in a previous study in the ventral cochlear nucleus (Pressnitzer et al., 2000). The main results were: (1) The shape of the responses of on-sustained units to ramped sinusoids resembled the shape of the responses to damped sinusoids. This is in contrast to the response shapes in the VCN, which were always similar to the stimulating sinusoid. (2) Units classified as onsets often responded only to the damped stimuli. (3) All units display significant asymmetry in discharge rate for at least one of the half-lives tested and 20% showed significant response asymmetry over all of the half-lives tested. (4) A summary population measure of temporal asymmetry based on total spike count reveals the same pattern of results as that obtained psychophysically using the same stimuli (Patterson, 1994a).

The lower limit of melodic pitch.

An objective melody task was used to determine the lower limit of melodic pitch (LLMP) for harmonic complex tones. The LLMP was defined operationally as the repetition rate below which listeners could no longer recognize that one of the notes in a four-note, chromatic melody had changed by a semitone. In the first experiment, the stimuli were broadband tones with all their components in cosine phase, and the LLMP was found to be around 30 Hz. In the second experiment, the tones were filtered into bands about 1 kHz in width to determine the influence of frequency region on the LLMP. The results showed that whenever there was energy present below 800 Hz, the LLMP was still around 30 Hz. When the energy was limited to higher-frequency regions, however, the LLMP increased progressively, up to 270 Hz when the energy was restricted to the region above 3.2 kHz. In the third experiment, the phase relationship between spectral components was altered to determine whether the shape of the waveform affects the LLMP. When the envelope peak factor was reduced using the Schroeder phase relationship, the LLMP was not affected. When a secondary peak was introduced into the envelope of the stimuli by alternating the phase of successive components between two fixed values, there was a substantial reduction in the LLMP, for stimuli containing low-frequency energy. A computational auditory model that extracts pitch information with autocorrelation can reproduce all of the observed effects, provided the contribution of longer time intervals is progressively reduced by a linear weighting function that limits the mechanism to time intervals of less than about 33 ms.

The responses of single units in the ventral cochlear nucleus of the guinea pig to damped and ramped sinusoids.

Human listeners hear an asymmetry in the perception of damped and ramped sinusoids; the partial loudness of the envelope component is greater than the partial loudness of the carrier component for damped sinusoids. Here we show that an asymmetry also occurs in the physiological responses of most units in the ventral cochlear nucleus to these same sounds. The activity elicited by damped sinusoids is mainly restricted to the beginning of each envelope period, which is not the case for ramped sinusoids. This can be quantified by computing the ratio of the tallest bin of the modulation period histogram to the total number of spikes (the peak-to-total ratio, p/t). Damped sinusoids produce a higher p/t than ramped sinusoids, which demonstrates physiological temporal asymmetry. It is also the case that ramped sinusoids typically elicit more spikes than damped sinusoids. The physiological asymmetry occurs where the perceptual asymmetry is present. It is maximal at modulation half-lives of 4 and 16 ms, greatly reduced at 1 ms and absent at 64 ms. Different unit types exhibit differing degrees of temporal asymmetry. Onset units produce the greatest p/t asymmetry, primary-like units produce the least asymmetry and chopper units are in-between. With regard to total spike count, the maximal asymmetry occurs with chopper units. If primary-like units are assumed to reflect the activity in primary auditory nerve fibres, then there is enhancement of temporal asymmetry in the ventral cochlear nucleus by both onset and chopper units.

The lower limit of pitch as determined by rate discrimination.

This paper is concerned with the lower limit of pitch for complex, harmonic sounds, like the notes produced by low-pitched musical instruments. The lower limit of pitch is investigated by measuring rate discrimination thresholds for harmonic tones filtered into 1.2-kHz-wide bands with a lower cutoff frequency, F(c), ranging from 0.2 to 6.4 kHz. When F(c) is below 1 kHz and the harmonics are in cosine phase, rate discrimination threshold exhibits a rapid, tenfold decrease as the repetition rate is increased from 16 to 64 Hz, and over this range, the perceptual quality of the stimuli changes from flutter to pitch. When F(c) is increased above 1 kHz, the slope of the transition from high to low thresholds becomes shallower and occurs at progressively higher rates. A quantitative comparison of the cosine-phase thresholds with subjective estimates of the existence region of pitch from the literature shows that the transition in rate discrimination occurs at approximately the same rate as the lower limit of pitch. The rate discrimination experiment was then repeated with alternating-phase harmonic tones whose envelopes repeat at twice the repetition rate of the waveform. In this case, when F(c) is below 1 kHz, the transition in rate discrimination is shifted downward by almost an octave relative to the transition in the cosine-phase thresholds. The results support the hypothesis that in the low-frequency region, the pitch limit is determined by a temporal mechanism, which analyzes time intervals between peaks in the neural activity pattern. It seems that temporal processing of pitch is limited to time intervals less than 33 ms, corresponding to a pitch limit of about 30 Hz.

Perception of musical tension for nontonal orchestral timbres and its relation to psychoacoustic roughness.

Can tension in nontonal music be expressed without dynamic or rhythmic cues? Perceptual theories of tonal harmony predict that psychoacoustic roughness plays an important role in the perception of this tension. We chose a set of orchestrated chords from a nontonal piece and investigated listeners' judgments of musical tension and roughness. Paired comparisons yielded psychophysical scales of tension and roughness. Two experiments established distinct levels of these two attributes across chords. A model simulation reproduced the experimental roughness measures. The results indicate that nontonal tension could be perceived consistently on the basis of timbral differences and that it was correlated with roughness, the correlation being stronger as the perceptual salience of other attributes (such as high-pitched tones or tonal intervals) was reduced.

Two phase effects in roughness perception.

The respective influences of spectral and temporal aspects of sound in roughness perception are examined by way of phase manipulations. In a first experiment, the phase of the central component of three-component signals is shown to modify perceived roughness, for a given amplitude spectrum, regardless of whether it modifies the waveform envelope. A second experiment shows that the shape of the waveform envelope, for a given amplitude spectrum and a given modulation depth, also influences perceived roughness. We interpret both of these results by considering the envelope of an internal representation that is deduced from the physical signal by taking into account peripheral auditory processing. The results indicate that the modulation depth of such an internal representation is not the only determinant of roughness, but that an effect of temporal asymmetry is also to be taken into account.