The Music of Silence: Part II: Music Listening Induces Imagery Responses.

During music listening, humans routinely acquire the regularities of the acoustic sequences and use them to anticipate and interpret the ongoing melody. Specifically, in line with this predictive framework, it is thought that brain responses during such listening reflect a comparison between the bottom-up sensory responses and top-down prediction signals generated by an internal model that embodies the music exposure and expectations of the listener. To attain a clear view of these predictive responses, previous work has eliminated the sensory inputs by inserting artificial silences (or sound omissions) that leave behind only the corresponding predictions of the thwarted expectations. Here, we demonstrate a new alternate approach in which we decode the predictive electroencephalography (EEG) responses to the silent intervals that are naturally interspersed within the music. We did this as participants (experiment 1, 20 participants, 10 female; experiment 2, 21 participants, 6 female) listened or imagined Bach piano melodies. Prediction signals were quantified and assessed via a computational model of the melodic structure of the music and were shown to exhibit the same response characteristics when measured during listening or imagining. These include an inverted polarity for both silence and imagined responses relative to listening, as well as response magnitude modulations that precisely reflect the expectations of notes and silences in both listening and imagery conditions. These findings therefore provide a unifying view that links results from many previous paradigms, including omission reactions and the expectation modulation of sensory responses, all in the context of naturalistic music listening.SIGNIFICANCE STATEMENT Music perception depends on our ability to learn and detect melodic structures. It has been suggested that our brain does so by actively predicting upcoming music notes, a process inducing instantaneous neural responses as the music confronts these expectations. Here, we studied this prediction process using EEGs recorded while participants listen to and imagine Bach melodies. Specifically, we examined neural signals during the ubiquitous musical pauses (or silent intervals) in a music stream and analyzed them in contrast to the imagery responses. We find that imagined predictive responses are routinely co-opted during ongoing music listening. These conclusions are revealed by a new paradigm using listening and imagery of naturalistic melodies.

The Music of Silence: Part I: Responses to Musical Imagery Encode Melodic Expectations and Acoustics.

Musical imagery is the voluntary internal hearing of music in the mind without the need for physical action or external stimulation. Numerous studies have already revealed brain areas activated during imagery. However, it remains unclear to what extent imagined music responses preserve the detailed temporal dynamics of the acoustic stimulus envelope and, crucially, whether melodic expectations play any role in modulating responses to imagined music, as they prominently do during listening. These modulations are important as they reflect aspects of the human musical experience, such as its acquisition, engagement, and enjoyment. This study explored the nature of these modulations in imagined music based on EEG recordings from 21 professional musicians (6 females and 15 males). Regression analyses were conducted to demonstrate that imagined neural signals can be predicted accurately, similarly to the listening task, and were sufficiently robust to allow for accurate identification of the imagined musical piece from the EEG. In doing so, our results indicate that imagery and listening tasks elicited an overlapping but distinctive topography of neural responses to sound acoustics, which is in line with previous fMRI literature. Melodic expectation, however, evoked very similar frontal spatial activation in both conditions, suggesting that they are supported by the same underlying mechanisms. Finally, neural responses induced by imagery exhibited a specific transformation from the listening condition, which primarily included a relative delay and a polarity inversion of the response. This transformation demonstrates the top-down predictive nature of the expectation mechanisms arising during both listening and imagery.SIGNIFICANCE STATEMENT It is well known that the human brain is activated during musical imagery: the act of voluntarily hearing music in our mind without external stimulation. It is unclear, however, what the temporal dynamics of this activation are, as well as what musical features are precisely encoded in the neural signals. This study uses an experimental paradigm with high temporal precision to record and analyze the cortical activity during musical imagery. This study reveals that neural signals encode music acoustics and melodic expectations during both listening and imagery. Crucially, it is also found that a simple mapping based on a time-shift and a polarity inversion could robustly describe the relationship between listening and imagery signals.

Accurate Decoding of Imagined and Heard Melodies.

Music perception requires the human brain to process a variety of acoustic and music-related properties. Recent research used encoding models to tease apart and study the various cortical contributors to music perception. To do so, such approaches study temporal response functions that summarise the neural activity over several minutes of data. Here we tested the possibility of assessing the neural processing of individual musical units (bars) with electroencephalography (EEG). We devised a decoding methodology based on a maximum correlation metric across EEG segments (maxCorr) and used it to decode melodies from EEG based on an experiment where professional musicians listened and imagined four Bach melodies multiple times. We demonstrate here that accurate decoding of melodies in single-subjects and at the level of individual musical units is possible, both from EEG signals recorded during listening and imagination. Furthermore, we find that greater decoding accuracies are measured for the maxCorr method than for an envelope reconstruction approach based on backward temporal response functions (bTRF env ). These results indicate that low-frequency neural signals encode information beyond note timing, especially with respect to low-frequency cortical signals below 1 Hz, which are shown to encode pitch-related information. Along with the theoretical implications of these results, we discuss the potential applications of this decoding methodology in the context of novel brain-computer interface solutions.

Learning Speech Production and Perception through Sensorimotor Interactions.

Action and perception are closely linked in many behaviors necessitating a close coordination between sensory and motor neural processes so as to achieve a well-integrated smoothly evolving task performance. To investigate the detailed nature of these sensorimotor interactions, and their role in learning and executing the skilled motor task of speaking, we analyzed ECoG recordings of responses in the high-γ band (70-150 Hz) in human subjects while they listened to, spoke, or silently articulated speech. We found elaborate spectrotemporally modulated neural activity projecting in both "forward" (motor-to-sensory) and "inverse" directions between the higher-auditory and motor cortical regions engaged during speaking. Furthermore, mathematical simulations demonstrate a key role for the forward projection in "learning" to control the vocal tract, beyond its commonly postulated predictive role during execution. These results therefore offer a broader view of the functional role of the ubiquitous forward projection as an important ingredient in learning, rather than just control, of skilled sensorimotor tasks.