An implicit representation of stimulus ambiguity in pupil size.

To guide behavior, perceptual systems must operate on intrinsically ambiguous sensory input. Observers are usually able to acknowledge the uncertainty of their perception, but in some cases, they critically fail to do so. Here, we show that a physiological correlate of ambiguity can be found in pupil dilation even when the observer is not aware of such ambiguity. We used a well-known auditory ambiguous stimulus, known as the tritone paradox, which can induce the perception of an upward or downward pitch shift within the same individual. In two experiments, behavioral responses showed that listeners could not explicitly access the ambiguity in this stimulus, even though their responses varied from trial to trial. However, pupil dilation was larger for the more ambiguous cases. The ambiguity of the stimulus for each listener was indexed by the entropy of behavioral responses, and this entropy was also a significant predictor of pupil size. In particular, entropy explained additional variation in pupil size independent of the explicit judgment of confidence in the specific situation that we investigated, in which the two measures were decoupled. Our data thus suggest that stimulus ambiguity is implicitly represented in the brain even without explicit awareness of this ambiguity.

Consonance Perception in Congenital Amusia: Behavioral and Brain Responses to Harmonicity and Beating Cues.

Congenital amusia is a neurodevelopmental disorder characterized by difficulties in the perception and production of music, including the perception of consonance and dissonance, or the judgment of certain combinations of pitches as more pleasant than others. Two perceptual cues for dissonance are inharmonicity (the lack of a common fundamental frequency between components) and beating (amplitude fluctuations produced by close, interacting frequency components). Amusic individuals have previously been reported to be insensitive to inharmonicity, but to exhibit normal sensitivity to beats. In the present study, we measured adaptive discrimination thresholds in amusic participants and found elevated thresholds for both cues. We recorded EEG and measured the MMN in evoked potentials to consonance and dissonance deviants in an oddball paradigm. The amplitude of the MMN response was similar overall for amusic and control participants; however, in controls, there was a tendency toward larger MMNs for inharmonicity than for beating cues, whereas the opposite tendency was observed for the amusic participants. These findings suggest that initial encoding of consonance cues may be intact in amusia despite impaired behavioral performance, but that the relative weight of nonspectral (beating) cues may be increased for amusic individuals.

Short- and long-term memory for pitch and non-pitch contours: Insights from congenital amusia.

Congenital amusia is a neurodevelopmental disorder characterized by deficits in music perception, including discriminating and remembering melodies and melodic contours. As non-amusic listeners can perceive contours in dimensions other than pitch, such as loudness and brightness, our present study investigated whether amusics' pitch contour deficits also extend to these other auditory dimensions. Amusic and control participants performed an identification task for ten familiar melodies and a short-term memory task requiring the discrimination of changes in the contour of novel four-tone melodies. For both tasks, melodic contour was defined by pitch, brightness, or loudness. Amusic participants showed some ability to extract contours in all three dimensions. For familiar melodies, amusic participants showed impairment in all conditions, perhaps reflecting the fact that the long-term memory representations of the familiar melodies were defined in pitch. In the contour discrimination task with novel melodies, amusic participants exhibited less impairment for loudness-based melodies than for pitch- or brightness-based melodies, suggesting some specificity of the deficit for spectral changes, if not for pitch alone. The results suggest pitch and brightness may not be processed by the same mechanisms as loudness, and that short-term memory for loudness contours may be spared to some degree in congenital amusia.

Pitch discrimination with mixtures of three concurrent harmonic complexes.

In natural listening contexts, especially in music, it is common to hear three or more simultaneous pitches, but few empirical or theoretical studies have addressed how this is achieved. Place and pattern-recognition theories of pitch require at least some harmonics to be spectrally resolved for pitch to be extracted, but it is unclear how often such conditions exist when multiple complex tones are presented together. In three behavioral experiments, mixtures of three concurrent complexes were filtered into a single bandpass spectral region, and the relationship between the fundamental frequencies and spectral region was varied in order to manipulate the extent to which harmonics were resolved either before or after mixing. In experiment 1, listeners discriminated major from minor triads (a difference of 1 semitone in one note of the triad). In experiments 2 and 3, listeners compared the pitch of a probe tone with that of a subsequent target, embedded within two other tones. All three experiments demonstrated above-chance performance, even in conditions where the combinations of harmonic components were unlikely to be resolved after mixing, suggesting that fully resolved harmonics may not be necessary to extract the pitch from multiple simultaneous complexes.

Auditory cortex and beyond: Deficits in congenital amusia.

Congenital amusia is a neuro-developmental disorder of music perception and production, with the observed deficits contrasting with the sophisticated music processing reported for the general population. Musical deficits within amusia have been hypothesized to arise from altered pitch processing, with impairments in pitch discrimination and, notably, short-term memory. We here review research investigating its behavioral and neural correlates, in particular the impairments at encoding, retention, and recollection of pitch information, as well as how these impairments extend to the processing of pitch cues in speech and emotion. The impairments have been related to altered brain responses in a distributed fronto-temporal network, which can be observed also at rest. Neuroimaging studies revealed changes in connectivity patterns within this network and beyond, shedding light on the brain dynamics underlying auditory cognition. Interestingly, some studies revealed spared implicit pitch processing in congenital amusia, showing the power of implicit cognition in the music domain. Building on these findings, together with audiovisual integration and other beneficial mechanisms, we outline perspectives for training and rehabilitation and the future directions of this research domain.

Familiar Tonal Context Improves Accuracy of Pitch Interval Perception.

A fundamental feature of everyday music perception is sensitivity to familiar tonal structures such as musical keys. Many studies have suggested that a tonal context can enhance the perception and representation of pitch. Most of these studies have measured response time, which may reflect expectancy as opposed to perceptual accuracy. We instead used a performance-based measure, comparing participants' ability to discriminate between a "small, in-tune" interval and a "large, mistuned" interval in conditions that involved familiar tonal relations (diatonic, or major, scale notes), unfamiliar tonal relations (whole-tone or mistuned-diatonic scale notes), repetition of a single pitch, or no tonal context. The context was established with a brief sequence of tones in Experiment 1 (melodic context), and a cadence-like two-chord progression in Experiment 2 (harmonic context). In both experiments, performance significantly differed across the context conditions, with a diatonic context providing a significant advantage over no context; however, no correlation with years of musical training was observed. The diatonic tonal context also provided an advantage over the whole-tone scale context condition in Experiment 1 (melodic context), and over the mistuned scale or repetition context conditions in Experiment 2 (harmonic context). However, the relatively small benefit to performance suggests that the main advantage of tonal context may be priming of expected stimuli, rather than enhanced accuracy of pitch interval representation.

Expectations for melodic contours transcend pitch.

The question of what makes a good melody has interested composers, music theorists, and psychologists alike. Many of the observed principles of good "melodic continuation" involve melodic contour-the pattern of rising and falling pitch within a sequence. Previous work has shown that contour perception can extend beyond pitch to other auditory dimensions, such as brightness and loudness. Here, we show that the generalization of contour perception to nontraditional dimensions also extends to melodic expectations. In the first experiment, subjective ratings for 3-tone sequences that vary in brightness or loudness conformed to the same general contour-based expectations as pitch sequences. In the second experiment, we modified the sequence of melody presentation such that melodies with the same beginning were blocked together. This change produced substantively different results, but the patterns of ratings remained similar across the 3 auditory dimensions. Taken together, these results suggest that (a) certain well-known principles of melodic expectation (such as the expectation for a reversal following a skip) are dependent on long-term context, and (b) these expectations are not unique to the dimension of pitch and may instead reflect more general principles of perceptual organization.