Neural correlates of musical timbre: an ALE meta-analysis of neuroimaging data.

Timbre is a central aspect of music that allows listeners to identify musical sounds and conveys musical emotion, but also allows for the recognition of actions and is an important structuring property of music. The former functions are known to be implemented in a ventral auditory stream in processing musical timbre. While the latter functions are commonly attributed to areas in a dorsal auditory processing stream in other musical domains, its involvement in musical timbre processing is so far unknown. To investigate if musical timbre processing involves both dorsal and ventral auditory pathways, we carried out an activation likelihood estimation (ALE) meta-analysis of 18 experiments from 17 published neuroimaging studies on musical timbre perception. We identified consistent activations in Brodmann areas (BA) 41, 42, and 22 in the bilateral transverse temporal gyri, the posterior superior temporal gyri and planum temporale, in BA 40 of the bilateral inferior parietal lobe, in BA 13 in the bilateral posterior Insula, and in BA 13 and 22 in the right anterior insula and superior temporal gyrus. The vast majority of the identified regions are associated with the dorsal and ventral auditory processing streams. We therefore propose to frame the processing of musical timbre in a dual-stream model. Moreover, the regions activated in processing timbre show similarities to the brain regions involved in processing several other fundamental aspects of music, indicating possible shared neural bases of musical timbre and other musical domains.

Lights on music cognition: A systematic and critical review of fNIRS applications and future perspectives.

Research investigating the neural processes related to music perception and production constitutes a well-established field within the cognitive neurosciences. While most neuroimaging tools have limitations in studying the complexity of musical experiences, functional Near-Infrared Spectroscopy (fNIRS) represents a promising, relatively new tool for studying music processes in both laboratory and ecological settings, which is also suitable for both typical and pathological populations across development. Here we systematically review fNIRS studies on music cognition, highlighting prospects and potentialities. We also include an overview of fNIRS basic theory, together with a brief comparison to characteristics of other neuroimaging tools. Fifty-nine studies meeting inclusion criteria (i.e., using fNIRS with music as the primary stimulus) are presented across five thematic sections. Critical discussion of methodology leads us to propose guidelines of good practices aiming for robust signal analyses and reproducibility. A continuously updated world map is proposed, including basic information from studies meeting the inclusion criteria. It provides an organized, accessible, and updatable reference database, which could serve as a catalyst for future collaborations within the community. In conclusion, fNIRS shows potential for investigating cognitive processes in music, particularly in ecological contexts and with special populations, aligning with current research priorities in music cognition.

Hierarchical syntax model of music predicts theta power during music listening.

Linguistic research showed that the depth of syntactic embedding is reflected in brain theta power. Here, we test whether this also extends to non-linguistic stimuli, specifically music. We used a hierarchical model of musical syntax to continuously quantify two types of expert-annotated harmonic dependencies throughout a piece of Western classical music: prolongation and preparation. Prolongations can roughly be understood as a musical analogue to linguistic coordination between constituents that share the same function (e.g., 'pizza' and 'pasta' in 'I ate pizza and pasta'). Preparation refers to the dependency between two harmonies whereby the first implies a resolution towards the second (e.g., dominant towards tonic; similar to how the adjective implies the presence of a noun in 'I like spicy … '). Source reconstructed MEG data of sixty-five participants listening to the musical piece was then analysed. We used Bayesian Mixed Effects models to predict theta envelope in the brain, using the number of open prolongation and preparation dependencies as predictors whilst controlling for audio envelope. We observed that prolongation and preparation both carry independent and distinguishable predictive value for theta band fluctuation in key linguistic areas such as the Angular, Superior Temporal, and Heschl's Gyri, or their right-lateralised homologues, with preparation showing additional predictive value for areas associated with the reward system and prediction. Musical expertise further mediated these effects in language-related brain areas. Results show that predictions of precisely formalised music-theoretical models are reflected in the brain activity of listeners which furthers our understanding of the perception and cognition of musical structure.

Inter-subject correlation of electroencephalographic and behavioural responses reflects time-varying engagement with natural music.

Musical engagement can be conceptualized through various activities, modes of listening and listener states. Recent research has reported that a state of focused engagement can be indexed by the inter-subject correlation (ISC) of audience responses to a shared naturalistic stimulus. While statistically significant ISC has been reported during music listening, we lack insight into the temporal dynamics of engagement over the course of musical works-such as those composed in the Western classical style-which involve the formulation of expectations that are realized or derailed at subsequent points of arrival. Here, we use the ISC of electroencephalographic (EEG) and continuous behavioural (CB) responses to investigate the time-varying dynamics of engagement with functional tonal music. From a sample of adult musicians who listened to a complete cello concerto movement, we found that ISC varied throughout the excerpt for both measures. In particular, significant EEG ISC was observed during periods of musical tension that built to climactic highpoints, while significant CB ISC corresponded more to declarative entrances and points of arrival. Moreover, we found that a control stimulus retaining envelope characteristics of the intact music, but little other temporal structure, also elicited significantly correlated EEG and CB responses, though to lesser extents than the original version. In sum, these findings shed light on the temporal dynamics of engagement during music listening and clarify specific aspects of musical engagement that may be indexed by each measure.

Recognizing structure in novel tunes: differences between human and rats.

A central feature in music is the hierarchical organization of its components. Musical pieces are not a simple concatenation of chords, but are characterized by rhythmic and harmonic structures. Here, we explore if sensitivity to music structure might emerge in the absence of any experience with musical stimuli. For this, we tested if rats detect the difference between structured and unstructured musical excerpts and compared their performance with that of humans. Structured melodies were excerpts of Mozart's sonatas. Unstructured melodies were created by the recombination of fragments of different sonatas. We trained listeners (both human participants and Long-Evans rats) with a set of structured and unstructured excerpts, and tested them with completely novel excerpts they had not heard before. After hundreds of training trials, rats were able to tell apart novel structured from unstructured melodies. Human listeners required only a few trials to reach better performance than rats. Interestingly, such performance was increased in humans when tonality changes were included, while it decreased to chance in rats. Our results suggest that, with enough training, rats might learn to discriminate acoustic differences differentiating hierarchical music structures from unstructured excerpts. More importantly, the results point toward species-specific adaptations on how tonality is processed.

Negative Space: An Alternative Framework for Archaeoacoustics.

Hearing the remote past seems impossible. Archaeoacoustics is a contemporary field intent on reconstructing the evolution of early communication systems, offering the possibility of developing methodologies relating to past sound signaling and music. Through a contribution of the emerging sensory field of archaeoacoustics and an example of acoustic assessments conducted at the site of Coves del Toll, can we understand signals of the past in order to investigate human behaviour and trace its cognitive evolution? This paper explores alternative methodological and theoretical approaches to understanding prehistoric sonic behaviours in early hominids and aims to set out a framework to theoretically and philosophically approach the "sound record" of the past. The theoretical proposition of this paper integrates the musical and sound art disciplines of spectralism and sound ecology to challenge the current limitations of listening to sound.

Prokofiev was (almost) right: A cross-cultural investigation of auditory-conceptual associations in Peter and the Wolf.

Over recent decades, studies investigating cross-modal correspondences have documented the existence of a wide range of consistent cross-modal associations between simple auditory and visual stimuli or dimensions (e.g., pitch-lightness). Far fewer studies have investigated the association between complex and realistic auditory stimuli and visually presented concepts (e.g., musical excerpts-animals). Surprisingly, however, there is little evidence concerning the extent to which these associations are shared across cultures. To address this gap in the literature, two experiments using a set of stimuli based on Prokofiev's symphonic fairy tale Peter and the Wolf are reported. In Experiment 1, 293 participants from several countries and with very different language backgrounds rated the association between the musical excerpts, images and words representing the story's characters (namely, bird, duck, wolf, cat, and grandfather). The results revealed that participants tended to consistently associate the wolf and the bird with the corresponding musical excerpt, while the stimuli of other characters were not consistently matched across participants. Remarkably, neither the participants' cultural background, nor their musical expertise affected the ratings. In Experiment 2, 104 participants were invited to rate each stimulus on eight emotional features. The results revealed that the emotional profiles associated with the music and with the concept of the wolf and the bird were perceived as more consistent between observers than the emotional profiles associated with the music and the concept of the duck, the cat, and the grandpa. Taken together, these findings therefore suggest that certain auditory-conceptual associations are perceived consistently across cultures and may be mediated by emotional associations.

Temporal speed prevails on interval duration in the SNARC-like effect for tempo.

The Spatial-Numerical Association of Response Codes (SNARC) effect is evidence of an association between number magnitude and response position, with faster left-key responses to small numbers and faster right-key responses to large numbers. Similarly, recent studies revealed a SNARC-like effect for tempo, defined as the speed of an auditory sequence, with faster left-key responses to slow tempo and faster right-key responses to fast tempo. In order to address some methodological issues of previous studies, in the present study we designed an experiment to investigate the occurrence of a SNARC-like effect for tempo, employing a novel procedure in which only two auditory beats in sequence with a very short interstimulus interval were used. In the "temporal speed" condition, participants were required to judge the temporal speed (slow or fast) of the sequence. In the "interval duration" condition, participants were required to judge the duration of the interval between the two beats (short or long). The results revealed a consistent SNARC-like effect in both conditions, with faster left-hand responses to slow tempo and faster right-hand responses to fast tempo. Interestingly, the consistency of the results across the two conditions indicates that the direction of the SNARC-like effect was influenced by temporal speed even when participants were explicitly required to focus on interval duration. Overall, the current study extends previous findings by employing a new paradigm that addresses potential confounding factors and strengthens evidence for the SNARC-like effect for tempo.

The development of rhythmic categories as revealed through an iterative production task.

Both humans and non-humans (e.g. birds and primates) preferentially produce and perceive auditory rhythms with simple integer ratios. In addition, these preferences (biases) tend to reflect specific integer-ratio rhythms that are common to one's cultural listening experience. To better understand the developmental trajectory of these biases, we estimated children's rhythm biases across the entire rhythm production space of simple (e.g., ratios of 1, 2, and 3) three-interval rhythms. North American children aged 6-11 years completed an iterative rhythm production task, in which they attempted to tap in synchrony with repeating three-interval rhythms chosen randomly from the space. For each rhythm, the child's produced rhythm was presented back to them as the stimulus, and over the course of 5 such iterations we used their final reproductions to estimate their rhythmic biases or priors. Results suggest that regardless of the initial rhythm, after 5 iterations, children's tapping converged on rhythms with (nearly) simple integer ratios, indicating that, like adults, their rhythmic priors consist of rhythms with simple-integer ratios. Furthermore, the relative weights (or prominence of different rhythmic priors) observed in children were highly correlated with those of adults. However, we also observed some age-related changes, especially for the ratio types that vary most across cultures. In an additional rhythm perception task, children were better at detecting rhythmic disruptions to a culturally familiar rhythm (in 4/4 m with 2:1:1 ratio pattern) than to a culturally unfamiliar rhythm (7/8 m with 3:2:2 ratios), and performance in this task was correlated with tapping variability in the iterative task. Taken together, our findings provide evidence that children as young as 6-years-old exhibit simple integer-ratio categorical rhythm priors in their rhythm production that closely resemble those of adults in the same culture.

An integrated review of music cognition and rhythmic stimuli in sensorimotor neurocognition and neurorehabilitation.

This work reviews the growing body of interdisciplinary research on music cognition, using biomechanical, kinesiological, clinical, psychosocial, and sociological methods. The review primarily examines the relationship between temporal elements in music and motor responses under varying contexts, with considerable relevance for clinical rehabilitation. After providing an overview of the terminology and approaches pertinent to theories of rhythm and meter from the musical-theoretical and cognitive fields, this review focuses on studies on the effects of rhythmic sensory stimulation on gait, rhythmic cues' effect on the motor system, reactions to rhythmic stimuli attempting to synchronize mobility (i.e., musical embodiment), and the application of rhythm for motor rehabilitation for individuals with Parkinson's disease, stroke, mild cognitive impairment, Alzheimer's disease, and other neurodegenerative or neurotraumatic diseases. This work ultimately bridges the gap between the musical-theoretical and cognitive science fields to facilitate innovative research in which each discipline informs the other.

The effects of rhythmic structure on tapping accuracy.

Prior investigations of simple rhythms in familiar time signatures have shown the importance of several mechanisms; notably, those related to metricization and grouping. But there has been limited study of complex rhythms, including those in unfamiliar time signatures, such as are found outside mainstream Western music. Here, we investigate how the structures of 91 rhythms with nonisochronous onsets (mostly complex, several in unfamiliar time signatures) influence the accuracy, velocity, and timing of taps made by participants attempting to synchronize with these onsets. The onsets were piano-tone cues sounded at a well-formed subset of isochronous cymbal pulses; the latter occurring every 234 ms. We modelled tapping at both the rhythm level and the pulse level; the latter provides insight into how rhythmic structure makes some cues easier to tap and why incorrect (uncued) taps may occur. In our models, we use a wide variety of quantifications of rhythmic features, several of which are novel and many of which are indicative of underlying mechanisms, strategies, or heuristics. The results show that, for these tricky rhythms, taps are disrupted by unfamiliar period lengths and are guided by crude encodings of each rhythm: the density of rhythmic cues, their circular mean and variance, and recognizing common small patterns and the approximate positions of groups of cues. These lossy encodings are often counterproductive for discriminating between cued and uncued pulses and are quite different to mechanisms-such as metricization and emphasizing group boundaries-thought to guide tapping behaviours in learned and familiar rhythms.

Expectation adaptation for rare cadences in music: Item order matters in repetition priming.

Humans make predictions about future events in many domains, including when they listen to music. Previous accounts of harmonic expectation in music have emphasised the role of implicit musical knowledge acquired in the long term through the mechanism of statistical learning. However, it is not known whether listeners can adapt their expectations for unusual harmonies in the short term through repetition priming, and whether the extent of any short-term adaptation depends on the unfolding statistical structure of the music. To explore these possibilities, we presented 150 participants with phrases from Bach chorales that ended with a cadence that was either a priori likely or unlikely based on the long-term statistical structure of the corpus of chorales. While holding the 50-50 incidence of likely vs. unlikely cadences constant, we manipulated the order in which these phrases were presented such that the local probability of hearing an unlikely cadence changed throughout the experiment. For each phrase, participants provided two judgements: (a) a prospective rating of how confident they were in their expectations for the cadence, and (b) a retrospective rating of how well the presented cadence matched their expectations. While confidence ratings increased over the course of the experiment, the rate of change decreased as the local probability of an unexpected cadence increased. Participants' expectations favoured likely cadences over unlikely cadences on average, but their expectation ratings for unlikely cadences increased at a faster rate over the course of the experiment than for likely cadences, particularly when the local probability of hearing an unlikely cadence was high. Thus, despite entrenched long-term statistics about cadences, listeners can indeed adapt to unusual musical harmonies and are sensitive to the local statistical structure of the musical environment. We suggest that this adaptation is an instance of Bayesian belief updating, a domain-general process that accounts for expectation adaptation in multiple domains.

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.

Probabilistic modelling of microtiming perception.

Music performances are rich in systematic temporal irregularities called "microtiming", too fine-grained to be notated in a musical score but important for musical expression and communication. Several studies have examined listeners' preference for rhythms varying in microtiming, but few have addressed precisely how microtiming is perceived, especially in terms of cognitive mechanisms, making the empirical evidence difficult to interpret. Here we provide evidence that microtiming perception can be simulated as a process of probabilistic prediction. Participants performed an XAB discrimination test, in which an archetypal popular drum rhythm was presented with different microtiming. The results indicate that listeners could implicitly discriminate the mean and variance of stimulus microtiming. Furthermore, their responses were effectively simulated by a Bayesian model of entrainment, using a distance function derived from its dynamic posterior estimate over phase. Wide individual differences in participant sensitivity to microtiming were predicted by a model parameter likened to noisy timekeeping processes in the brain. Overall, this suggests that the cognitive mechanisms underlying perception of microtiming reflect a continuous inferential process, potentially driving qualitative judgements of rhythmic feel.

Absolute pitch judgments of familiar melodies generalize across timbre and octave.

Most listeners can determine when a familiar recording of music has been shifted in musical key by as little as one semitone (e.g., from B to C major). These findings appear to suggest that absolute pitch memory is widespread in the general population. However, the use of familiar recordings makes it unclear whether these findings genuinely reflect absolute melody-key associations for at least two reasons. First, listeners may be able to use spectral cues from the familiar instrumentation of the recordings to determine when a familiar recording has been shifted in pitch. Second, listeners may be able to rely solely on pitch height cues (e.g., relying on a feeling that an incorrect recording sounds "too high" or "too low"). Neither of these strategies would require an understanding of pitch chroma or musical key. The present experiments thus assessed whether listeners could make accurate absolute melody-key judgments when listening to novel versions of these melodies, differing from the iconic recording in timbre (Experiment 1) or timbre and octave (Experiment 2). Listeners in both experiments were able to select the correct-key version of the familiar melody at rates that were well above chance. These results fit within a growing body of research supporting the idea that most listeners, regardless of formal musical training, have robust representations of absolute pitch - based on pitch chroma - that generalize to novel listening situations. Implications for theories of auditory pitch memory are discussed.

Language-to-music transfer effects depend on the tone language: Akan vs. East Asian tone languages.

Recent research suggests that speaking a tone language confers benefits in processing pitch in nonlinguistic contexts such as music. This research largely compares speakers of nontone European languages (English, French) with speakers of tone languages in East Asia (Mandarin, Cantonese, Vietnamese, Thai). However, tone languages exist on multiple continents-notably, languages indigenous to Africa and the Americas. With one exception (Bradley, Psychomusicology, 26(4), 337-345, 2016), no research has assessed whether these tone languages also confer pitch processing advantages. Two studies presented a melody change detection task, using quasirandom note sequences drawn from Western major scale tone probabilities. Listeners were speakers of Akan, a tone language of Ghana, plus speakers from previously tested populations (nontone language speakers and East Asian tone language speakers). In both cases, East Asian tone language speakers showed the strongest musical pitch processing, but Akan speakers did not exceed nontone speakers, despite comparable or better instrument change detection. Results suggest more nuanced effects of tone languages on pitch processing. Greater numbers of tones, presence of contour tones in a language's tone inventory, or possibly greater functional load of tone may be more likely to confer pitch processing benefits than mere presence of tone contrasts.

Confronting ethical and social issues related to the genetics of musicality.

New interdisciplinary research into genetic influences on musicality raises a number of ethical and social issues for future avenues of research and public engagement. The historical intersection of music cognition and eugenics heightens the need to vigilantly weigh the potential risks and benefits of these studies and the use of their outcomes. Here, we bring together diverse disciplinary expertise (complex trait genetics, music cognition, musicology, bioethics, developmental psychology, and neuroscience) to interpret and guide the ethical use of findings from recent and future studies. We discuss a framework for incorporating principles of ethically and socially responsible conduct of musicality genetics research into each stage of the research lifecycle: study design, study implementation, potential applications, and communication.

Music in the Middle: A Culture-Cognition-Mediator Model of Musical Functionality.

Music is both universal, appearing in every known human culture, and culture-specific, often defying intelligibility across cultural boundaries. This duality has been the source of debate within the broad community of music researchers, and there have been significant disagreements both on the ontology of music as an object of study and the appropriate epistemology for that study. To help resolve this tension, I present a culture-cognition-mediator model that situates music as a mediator in the mutually constitutive cycle of cultures and selves representing the ways individuals both shape and are shaped by their cultural environments. This model draws on concepts of musical grammars and schema, contemporary theories in developmental and cultural psychology that blur the distinction between nature and nurture, and recent advances in cognitive neuroscience. Existing evidence of both directions of causality is presented, providing empirical support for the conceptual model. The epistemological consequences of this model are discussed, specifically with respect to transdisciplinarity, hybrid research methods, and several potential empirical applications and testable predictions as well as its import for broader ontological conversations around the evolutionary origins of music itself.

Incidental learning in music reading: The music contingency learning task.

The present report investigated whether nonmusicians can incidentally learn musical skills needed for sight-reading. On each trial, participants identified a note name written inside of a note on the musical staff. In Experiment 1, each note was presented frequently with the congruent note name (e.g., "do" with the note for "do") and rarely with the incongruent names (e.g., "do" with the note for "fa"). With or without deliberate learning instructions, a robust contingency learning effect was observed: faster responses for congruent trials compared with incongruent trials. Participants also explicitly identified the meaning of the note positions more accurately than chance. Experiment 2 ruled out the potential influence of preexisting knowledge on the contingency learning effect by presenting notes most often with an incongruent note name. Robust learning was again observed, suggesting that participants acquired sufficient knowledge of musical notation to produce automatic influences on behaviour (e.g., akin to the interference effect previously found in skilled musicians). A congruency effect was additionally observed in Experiment 2, however. Experiment 3 further explored to what extent this congruency effect might be due to prior music knowledge and/or spatial stimulus-response compatibility between note and response locations (analogous to the SMARC effect). Overall, our results open up new avenues for investigating the incidental learning of complex material, musical or otherwise, and for reinforcing learning even further.

Detection of pitch errors in well-known songs.

We examined pitch-error detection in well-known songs sung with or without meaningful lyrics. In Experiment 1, adults heard the initial phrase of familiar songs sung with lyrics or repeating syllables (la) and judged whether they heard an out-of-tune note. Half of the renditions had a single pitch error (50 or 100 cents); half were in tune. Listeners were poorer at pitch-error detection in songs with lyrics. In Experiment 2, within-note pitch fluctuations in the same performances were eliminated by auto-tuning. Again, pitch-error detection was worse for renditions with lyrics (50 cents), suggesting adverse effects of semantic processing. In Experiment 3, songs were sung with repeating syllables or scat syllables to ascertain the role of phonetic variability. Performance was poorer for scat than for repeating syllables, indicating adverse effects of phonetic variability, but overall performance exceeded Experiment 1. In Experiment 4, listeners evaluated songs in all styles (repeating syllables, scat, lyrics) within the same session. Performance was best with repeating syllables (50 cents) and did not differ between scat or lyric versions. In short, tracking the pitches of highly familiar songs was impaired by the presence of words, an impairment stemming primarily from phonetic variability rather than interference from semantic processing.