TABLE 1.

The motor system’s predictive role in music and rhythm perception is only one component of larger networks of sensorimotor processing, namely the dorsal auditory pathway and the motor cortico-basal ganglia-thalamo-cortical circuit.

Brain area	Authors	Proposed role of each brain area	Experimental task and stimulus type	Type of data
Cerebellum	Ivry and Schlerf, 2008	Dedicated timing mechanism; coordination of movement, internal timing mechanisms involved with sub-second timing	Theoretical Paper/Review
	Bastian, 2006	Predictive models of movement	Theoretical Paper/Review
	Nozaradan et al., 2017	Tracking beats in rhythms with fast tempos; more prominent role in absolute timing vs. relative timing	Passive Listening. Auditory rhythms designed to induce a beat – syncopated and unsyncopated.	EEG
	Gordon et al., 2018	Meta-analysis of fMRI studies of recruitment of motor system during music listening	Meta-analysis. Various listening tasks – auditory rhythms or music.	fMRI
Basal Ganglia	Merchant et al., 2013	Interacts with the cortico-thalamic-striatal circuit in a context dependent manner	Theoretical Paper/Review
	Coull and Nobre, 2008	Perceptual temporal expectation; explicit timing	Theoretical Paper/Review
	Nozaradan et al., 2017	Tracking beats in complex rhythm sequences	Passive Listening Auditory rhythms designed to induce a beat – syncopated and unsyncopated.	EEG
	Grahn, 2009	Internal beat generation; more prominent role in relative vs. absolute timing	Discrimination task, same or different judgment of two auditory stimuli. Auditory rhythms – beat-based structure and non-beat-based structure; Accents- duration or volume accented (externally generated) or unaccented (internally generated) beats.	fMRI/Behavioral
	Grahn et al., 2011	Internal representation of auditory rhythms that support cross-modal interactions in beat perception and generation	Discrimination task, rhythmic tempo change. Auditory tone metronome and visual flashing metronome. Two groups: one with auditory first visual second, and the other vice versa.	fMRI/Behavioral
	Grahn and Rowe, 2009	Internal beat generation: part of cortico-subcortical network involved in beat perception and generation	Indicate the strength of the perceived beat. Auditory rhythms of varying complexity and some with volume accents.	fMRI/Behavioral
	Grahn and Rowe, 2013	Putamen activity in beat prediction, but not beat finding	Attentive listening; occasionally indicate level of feeling of the beat. Auditory rhythms of varying intervals and rates, beat and non-beat (jittered) rhythms.	fMRI/Behavioral
	Grahn and Brett, 2007	Higher activity for rhythms with integer ratio relationships between intervals and with regular perceptual accents	1st experiment (behavioral) reproduce auditory rhythms. 2nd experiment (fMRI) indicate if the rhythm played matched previous rhythms. Metered auditory rhythms of varying integer intervals and complexity.	fMRI/Behavioral
	Teki et al., 2011	Striato-thalamo-cortical network involved in beat-based timing, while an olivocerebellar network involved in duration-based timing	Judge duration matches in a set of tones. Auditory tones, either isochronous or jittered, arranged in either rhythm-based or absolute duration-based sets.	fMRI/Behavioral
	Araneda et al., 2017	Hearing, feeling or seeing a beat recruits a supramodal network in the auditory dorsal stream	Discrimination task, between beat and non-beat rhythms. Auditory, visual, and vibrotactile rhythms.	fMRI/Behavioral
Primary and premotor cortices	Kilavik et al., 2014	Movement preparation, cue anticipation	Theoretical Paper/Review
	Schubotz, 2007	Predictive processing of external events, even in the absence of proprioceptive or interoceptive information	Theoretical Paper/Review
	Morillon and Baillet, 2017	Beta and delta oscillations directed to auditory cortex encode temporal predictions	Passive listening (listen condition); active tapping with the beat (tracking condition). Auditory melody – different tones either on beat, anti-phase, or quasi-phase with the beat.	MEG/Behavioral
	Gordon et al., 2018	Meta-analysis of fMRI studies of recruitment of motor system during music listening	Meta-analysis. Various listening tasks – Auditory rhythms or music.	fMRI
Premotor cortex	Grahn and Rowe, 2009	Cortico-cortical coupling with SMA and auditory cortex in duration beat perception; part of cortico-subcortical network involved in beat perception and generation	Indicate the strength of the perceived beat. Auditory rhythms of varying complexity and some with volume accents.	fMRI/Behavioral
	Teki et al., 2011	Striato-thalamo-cortical network involved in beat-based timing, while an olivocerebellar network involved in duration based timing	Judge duration matches in a set of tones. Auditory tones, either isochronous or jittered, arranged in either rhythm-based or absolute duration-based sets.	fMRI/Behavioral
	Chen et al., 2008a	Motor regions recruited while listening to music rhythms	Experiment 1: Listen to rhythm passively then tap along with rhythm. Experiment 2: Listen to rhythm passively then tap along to rhythm without foreknowledge of being asked to tap with the rhythm auditory tones in simple, complex, or ambiguous rhythms.	fMRI/Behavioral
Supplementary motor area	Coull et al., 2016b	Perceptual and motor timing; Comparing the duration of perceptual events, error monitoring	Theoretical Paper/Review
	Ross et al., 2018b	Not causally implicated in perceptual auditory interval timing	Discrimination task – same/different judgment of auditory intervals; detection task – identification of tempo or phase shifted metronome click. Auditory intervals of pairs of tones; metronome click track over musical stimuli.	Behavioral (pre/post TMS down-regulatory stimulation)
	Grahn and Brett, 2007	Higher activity for rhythms with integer ratio relationships between intervals and with regular perceptual accents; in musicians: higher activity for all rhythms when compared to rest	Experiment 1 (behavioral): reproduce auditory rhythms. Experiment 2 (fMRI): indicate if the rhythm played matched previous rhythms. Metered auditory rhythms of varying integer intervals and complexity.	fMRI/Behavioral
	Grahn and McAuley, 2009	Stronger activity in strong beat-perceivers vs. weak beat-perceivers, no correlation with musicianship	Discrimination task, rhythmic tempo change. Auditory isochronous rhythms.	fMRI/Behavioral
	Grahn and Rowe, 2009	Coupling with STG in beat perception for musicians; part of cortico-subcortical network involved in beat perception and generation	Indicate the strength of the perceived beat. Auditory rhythms of varying complexity and some with volume accents.	fMRI/Behavioral
	Teki et al., 2011	Striato-thalamo-cortical network involved in beat-based time, while an olivocerebellar network involved in duration-based timing	Judge duration matches in a set of tones. Auditory tones, either isochronous or jittered, arranged in either rhythm-based or absolute duration-based sets.	fMRI/Behavioral
	Chen et al., 2008a	Motor regions recruited while listening to music rhythms	Experiment 1: Listen to rhythm passively then tap along with rhythm. Experiment 2: Listen to rhythm passively then tap along to rhythm without foreknowledge of being asked to tap with the rhythm auditory tones in simple, complex, or ambiguous rhythms.	fMRI/Behavioral
	Araneda et al., 2017	Hearing, feeling or seeing a beat recruits a supramodal network in the auditory dorsal stream	Discrimination task, between beat and non-beat rhythms. Auditory, visual, and vibrotactile rhythms.	fMRI/Behavioral
Medial premotor cortex	Merchant et al., 2014	Absolute and relative timing mechanisms within two separate neural populations	Theoretical Paper/Review
	Crowe et al., 2014	Absolute and relative timing mechanisms within two separate neural populations	Synchronization Continuation Task. Isochronous visual stimuli or auditory tones.	Behavioral; Extracellular activity of single neurons (in Macaca mulatta)
	Grahn and McAuley, 2009	Stronger activity in strong beat-perceivers vs. weak beat-perceivers, no correlation with musicianship	Discrimination task, rhythmic tempo change. Auditory isochronous rhythms.	fMRI/Behavioral
Parietal Cortex	Rauschecker, 2011; Merchant and Honing, 2014; Patel and Iversen, 2014	Interface between motor and auditory cortices, sensorimotor integration	Theoretical Papers/Reviews
	Coull and Nobre, 2008	Perceptual temporal expectation; implicit timing	Theoretical Paper/Review
	Coull et al., 2016a	Temporal predictability via fixed or dynamic predictions	Cued reaction time task. Visual cue that predicted target presentation time (temporal condition), or provided no information for target presentation (neutral condition) with variable intervals between cue and target.	fMRI/Behavioral
	Ross et al., 2018b	Causally implicated in perceptual beat-based timing	Discrimination task – same/different judgment of auditory intervals; detection task – identification of tempo or phase shifted metronome click. Auditory intervals of pairs of tones; metronome click track over musical stimuli.	Behavioral (pre/post TMS down-regulatory stimulation)
Auditory Cortex	Koelsch et al., 2019	Event related potentials associated with predictive processes in music	Theoretical Paper/Review
	Fujioka et al., 2012	Beta-band activity predicts onset of beats in music	Passive listening, while watching silent videos. Auditory isochronous rhythms of several tempos and one irregular rhythm.	MEG
	Fujioka et al., 2015	Beta-band activity represents timing information being translated for auditory-motor coordination	Passive listening to metered rhythms, followed by attentive listening to un-metered rhythms that the participants were asked to imagine as metered. March and Waltz metered rhythms	MEG
	Auksztulewicz et al., 2019	Temporal prediction of rhythm and beats	Identify target chords. Auditory rhythmic or jittered sequences of distractor chords preceding target chords.	MEG/EEG/Behavioral
	Honing et al., 2018	Event related potentials to perceptual deviants in rhythmic stimuli	Passive listening. Auditory oddball paradigm with isochronous or jittered rhythms.	EEG (of Macaca mulatta)
	Bouwer et al., 2016	Event related potentials to perceptual deviants in rhythmic stimuli; ERPs modulated by attention in musicians	Passive or attentive listening. Auditory oddball paradigm with isochronous or jittered rhythms.	EEG
Dopaminergic System/Striatal Dopamine	Schultz, 2007	Multiple time courses of dopamine changes mediate multiple time courses of behavioral processes	Theoretical Paper/Review
	Friston et al., 2009	Reward learning, encoding of precision	Theoretical Paper/Review
	Friston et al., 2012; FitzGerald et al., 2015	Reward learning, encoding of precision	Theoretical Papers/Computational Models	Simulated dopaminergic responses
	Sarno et al., 2017	Temporal expectation of perceptual cues; reward prediction error and (un)certainty	Detect weak vibrotactile stimuli. Variable interval durations between tactile start cue and vibrotactile stimuli.	Intracellular recording, monkey brain
	Koshimori et al., 2019	Rhythmic auditory stimulation (RAS) attenuates dopaminergic response	Synchronization task, RAS and no-RAS conditions; various auditory rhythms, single auditory beats or metronome clicks over instrumental music.	Behavioral/MRI/PET
	Brodal et al., 2017	Rhythmic music reduces connectivity between basal ganglia and reward system	Passive listening. Electronic dance music in a continuous-stimulation design.	fMRI

This table provides an overview of the brain areas and mechanisms which make up these networks and are involved in the predictive processing of auditory beat and rhythm perception. Each brain area is introduced with one or more Theoretical or Review Papers contextualizing that brain area’s proposed role, followed by a non-exhaustive list of supporting experimental work. This table is intended to serve as a tool for new or continuing researchers engaging in work on rhythm and musical beat perception.