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Pupil response reflects processing of polyrhythm and microtiming in musical grooves: A study of musicians and non- musicians Jo Fougner Skaansar Submitted as cand.psychol. thesis Department of Psychology UNIVERSITY OF OSLO October 2017

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Page 1: Pupil response reflects processing of polyrhythm and microtiming in musical grooves… · 2018. 1. 24. · Experiment 2, participants were exposed to original recordings of double

Pupil response reflects processing of

polyrhythm and microtiming in musical

grooves: A study of musicians and non-

musicians

Jo Fougner Skaansar

Submitted as cand.psychol. thesis

Department of Psychology

UNIVERSITY OF OSLO

October 2017

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Pupil response reflects processing of polyrhythm and microtiming in musical grooves: A study of musicians and non-musicians

By Jo Fougner Skaansar

Submitted as cand.psychol. thesis

Department of Psychology

UNIVERSITY OF OSLO

October 2017

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© Jo Fougner Skaansar

2017

Pupil response reflects processing of polyrhythm and microtiming in musical grooves:

A study of musicians and non-musicians

Author: Jo Fougner Skaansar

Supervisors: Bruno Laeng & Anne Danielsen

http://www.duo.uio.no

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Abstract Author: Jo Fougner Skaansar. Title: Pupil response reflects processing of polyrhythm and

microtiming in musical grooves: A study of musicians and non-musicians. Supervisors:

Bruno Laeng & Anne Danielsen.

Background and objectives: Instances of polyrhythm (cf. Vuust et al., 2006) and

microtiming (cf. Danielsen, Jensenius, & Haugen, 2015) appear commonly in groove-based

music, where they may challenge the listener’s internal framework of timing regularities,

often referred to as meter. Two influential theories, The Dynamic Attending Theory (DAT;

Jones & Boltz, 1989) and Predictive Coding Theory (PC; Vuust & Witek, 2014, cf. Friston

2005) hypothesize that attentional effort is recruited when metrical frameworks are

challenged. The present study addresses this hypothesis by asking: During listening to

musical groove-excerpts, are instances of polyrhythm and microtiming asynchrony related to

an increase in ‘mental effort’ (as indexed by pupillometry; Kahneman, 1973), as well as a

decrease in quality of sensorimotor synchronisation (as indexed by reduced finger tapping

accuracy)? In addition, the common assumption that microtiming promotes groove

experience is investigated. Of special interest is looking at effects of musical expertise on the

processing of polyrhythmic and microtiming events.

Method: Two experiments were designed by the author with help from the

supervisors. Data collection was done by the author. In Experiment 1, professional jazz

musicians (N = 16) and non-musicians (N = 16), matched demographically, were exposed to

a groove-based musical excerpt with a 4 against 3 polyrhythmic event that was contrasted

with a similar but non-polyrhythmic excerpt (stimuli borrowed from Vuust et al., 2006). In

Experiment 2, participants were exposed to original recordings of double bass and drum kit-

grooves of varying structural complexity, manipulated into five distinct microtiming

bass/drums-asynchrony conditions (-80 ms < X < 80 ms). All musical stimuli were presented

in a passive condition (‘listening only’) and an active tapping condition (‘synchronising with

the beat’). We recorded pupil diameter sizes and participants gave 1) their subjective ratings

after listening to each clip in the passive condition, and 2) tapping responses during the active

condition by pressing a key on the keyboard of a PC.

Main results: In Experiment 1, as expected, exposure to polyrhythm was related to

larger pupil sizes (more effort) and lower tapping accuracy compared to the control

condition. In Experiment 2, magnitudes of bidirectional bass/drums-microtiming

asynchronies were positively related to pupil dilation and negatively related to tapping

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accuracy. In both experiments, tapping the beat while listening yielded higher

psychophysiological effects than when listening only. Thus, the main effects of polyrhythm

and microtiming on pupil response and tapping accuracy supported both DAT and PC

accounts. Neither instances of polyrhythm or microtiming generated significant differences in

pupil response between musicians and non-musicians. However, professional jazz musicians

consistently showed superior tapping accuracy compared to non-musicians, which reflects

their enhanced expertise in rhythm perception and performance. On subjective ratings of

groove, participants generally preferred the on-the-grid grooves more than the grooves with

microtiming. Musicians showed a greater response than non-musicians on these ratings,

demonstrating their enhanced sensitivity to microtiming features in musical groove contexts.

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Acknowledgements For quite a few years I have been a psychology student during the day time and a jazz

musician at night. By doing a research project in music cognition, leading to this thesis, I had

a great opportunity to merge my two areas of interest. Throughout my musical career, timing,

microtiming, polyrhythm and groove, are all concepts I have had an intense, but still rather

tacit relationship with for many years. Thanks to outstanding support from my supervisors

Bruno Laeng (Department of Psychology, ‘RITMO Centre for Interdisciplinary Studies of

Rhythm, Time and Motion’) and Anne Danielsen (Department of Musicology, ‘RITMO

Centre for Interdisciplinary Studies of Rhythm, Time and Motion’), I was able to take a deep

dive into psychological aspects of rhythm and translate my musical and academic curiosity

into intriguing research questions, experimental design and, eventually, some very interesting

results. I owe both my supervisors great thanks; this project would not have been possible

without their extraordinary help and enthusiasm. Thanks to Bruno and Anne, I was also

fortunate to present a poster based on this thesis at the ‘Conference on Music & Eye-

Tracking’ that took part at the Max Planck Institut für Empirische Ästhetik, Frankfurt, 16-19

August, 2017.

I want to thank the participants who offered their time for participating in this study. I

would also like to express gratitude to my colleague and friend Jakop Filip Janssønn Hauan

for playing the drums on my microtiming stimuli, Martin Torvik Langerød at the Department

of Musicology for engineering the double bass recordings and preparing the experimental

sound clips, Kristian Nymoen at the Departments of Musicology/Informatics for preparation

of tapping data with custom MATLAB analyses, Gorm Helfjord for graphic aid on some of

the figures, and Rebekah Oomen for language consultation. Moreover, Dag-Erik Eilertsen at

the Department of Psychology, University of Oslo, as well as Peter Vuust and Maria Witek at

the ‘Center for Music in The Brain’ at Aarhus University, Denmark, gave me some valuable

ideas and comments on the project. A last thank you goes to my family, friends and Hedda

for invaluable support throughout this year.

The following are credited in the text with their initials:

-Anne Danielsen (AD)

-Martin Torvik Langerød (MTL)

-Kristian Nymoen (KN)

-Jakop Filip Janssønn Hauan (JJH)

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Table of Contents 1 Introduction.............................................................................................................................12 Theoreticalfoundation........................................................................................................42.1 Musicperceptionisenabledbyenculturationandrefinedbymusicaltraining..42.2 Perceptionofrhythm,beatandmeter................................................................................52.2.1 MeterperceptionI:MetricEncodingTheoryandDynamicAttendingTheory.............62.2.2 MeterperceptionII:PredictiveCodingTheory..........................................................................82.2.3 Developmentandneuralbasisofbeatandmeterperception.............................................8

2.3 Synchronisingtomusicalrhythm.........................................................................................92.4 Groove.........................................................................................................................................102.4.1 Twostructuralpropertiesofgroove-basedmusic.................................................................11Polyrhythm...........................................................................................................................................................................12Microtiming..........................................................................................................................................................................13

2.5 Pupillometry:ameasureofcognitiveprocessing........................................................152.5.1 Mentaleffortinthestudyofrhythmandmeterperception..............................................17

3 Thepresentstudy...............................................................................................................183.1 Hypothesesandpredictions................................................................................................183.2 Methods.......................................................................................................................................203.2.1 Participants..............................................................................................................................................20MusicalEarTest(MET)...................................................................................................................................................21

3.2.2 Stimuli........................................................................................................................................................22Experiment1:ThePolyrhythmExperiment..........................................................................................................22Experiment2:TheMicrotimingExperiment.........................................................................................................22

3.2.3 Setupandprocedure...........................................................................................................................243.2.4 Dataprocessing......................................................................................................................................27

4 Results....................................................................................................................................294.1 MusicEngagementQuestionnaire.....................................................................................294.2 Ageneralpupillarydriftingeffect......................................................................................294.3 Experiment1:ThePolyrhythmExperiment..................................................................304.3.1 Pupilresponses......................................................................................................................................304.3.2 Tappingaccuracy..................................................................................................................................30

4.4 Experiment2:TheMicrotimingExperiment..................................................................314.4.1 Pupilresponses......................................................................................................................................314.4.2 Tappingaccuracy..................................................................................................................................334.4.3 Ratingscales............................................................................................................................................35

5 GeneralDiscussion.............................................................................................................385.1 GeneraleffectsonpupilresponsesandtappingaccuracysupportDATandPC385.2 Effectsofmusicalexpertise..................................................................................................395.2.1 Pupillaryresponsewasunaffectedbymusicalexpertise....................................................405.2.2 Musiciansoutperformednon-musiciansintappingaccuracy...........................................42

5.3 Microtimingandtheexperienceofgroove.....................................................................435.4 Grooveexperience,effortandcomplexity......................................................................455.5 Limitationsandmethodologicalconsiderations..........................................................485.6 Futurestudies...........................................................................................................................49

6 Conclusion.............................................................................................................................507 Bibliography.........................................................................................................................517.1 References..................................................................................................................................517.2 Discography...............................................................................................................................63

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7.3 Filmography..............................................................................................................................648 Appendix................................................................................................................................688.1 FiguresA1-A7............................................................................................................................688.2 TablesA1-A5.............................................................................................................................71

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1 Introduction Music is a significant characteristic of human beings (Brown, 2004), found in all cultures

today and throughout history (Blacking, Byron, & Nettl, 1995). Listening to music may

enrich us with strong aesthetic experiences and emotions, recall of pleasant memories and the

urge to move our bodies. Music psychology traces back to the end of the 19th century and is

the fast-growing interdisciplinary

scientific field studying brain processes,

subjective experience and behaviour in

relation to the vibrations and sounds that

are categorized as music (Bonde, 2009).

Covering the field’s depth and width,

The Oxford Handbook of Music

Psychology (Hallam, Cross, & Thaut,

2016) presents theory and empirical

findings on cognitive perspectives,

aesthetics and emotional responses to

music, performance, improvisation,

music therapy and music education, to

name a few. Musicologists,

psychologists, philosophers,

anthropologists and computer scientists are among the scholars involved today in music

psychological research (Figure 1).

Music principally consists of melody, harmony, timbre/colour and rhythm. The

present study investigates the perception of rhythm; specifically, by placing under scrutiny

two rhythmic aspects, namely polyrhythm and microtiming. Polyrhythms can be defined as

(at least) two metric reference structures elapsing simultaneously within a single rhythmic

texture (Danielsen, 2006; Vuust, Roepstorff, Wallentin, Mouridsen, & Østergaard, 2006).

Microtiming refers to subtle timing asynchronies in music (Danielsen, 2015; Keil & Feld,

1994; Kilchenmann & Senn, 2015). Polyrhythm and microtiming are commonly occurring in

groove-based genres, including rock, jazz and hip-hop (Roholt, 2014), funk (Danielsen,

2006) and electronic dance music (EDM; Butler, 2006). Grooves are repetitive musical

patternsthat have a danceable quality to them. Moreover, they often contain certain features

of structural complexity (Pressing, 2002; Witek, 2017) that are assumed to contribute to a

Figure 1. Scope of the scientific discipline of music psychology. From Eagle (1996).

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groove experience. A groove experience refers to our urge to move our bodies and

synchronise with the music (Davies, Madison, Silva, & Gouyon, 2013), as well as pleasant

affect (Janata, Tomic, & Haberman, 2012).

Rhythm is experienced in the interaction of sounding events and underlying,

organizing and dynamic mental models (Danielsen, Jensenius, & Haugen, 2015). Theoretical

accounts of the relations between rhythmic input and temporal cognitive structures have been

addressed by Dynamic Attending Theory (DAT; Jones, 2016; Large & Jones, 1999; Large &

Snyder, 2009) and Predictive Coding Theory (PC; Vuust & Witek, 2014). The rhythmic

complexity added by polyrhythm and microtiming features may challenge listeners’

fundamental cognitive structures for musical timing (meter), a process that according to DAT

and PC demands attentional resources (effort; Danielsen et al., 2015; Vuust, Gebauer, &

Witek, 2014). Hence, the main objective of the present study, is to empirically test the

following theoretical prediction derived by DAT- and PC-accounts: Listening exposure to

rhythmic features (i.e., polyrhythm and microtiming asynchrony) in musical groove contexts

that challenge listeners’ metric framework, is 1) positively related to a psychophysiological

measure of mental effort like pupillometry, and 2) negatively related to a behavioural

measure of sensorimotor synchronisation quality, like finger tapping accuracy. According to

several cognitive psychologists, effort reflects the intensity of processing demands in the

brain (Just, Carpenter, & Miyake, 2003; Kahneman, 2011) and is often measured by phasic

pupil size diameter changes with the psychophysiological method of pupillometry, commonly

based on eye-tracking technology (Laeng, Sirois, & Gredeback, 2012). Finger-tapping

paradigms remain popular in investigating mechanisms of sensorimotor synchronisation

(Repp, 2005; Repp & Su, 2013). Sensorimotor synchronisation refers to ‘the coordination of

rhythmic movement with an external rhythm’ (Repp & Su, 2013, p. 403). Fluctuations in

intra-individual tapping accuracy have been found to reflect variation of external rhythmic

complexity (Chen, Penhune, & Zatorre, 2008), while inter-individual tapping accuracy

differences may reflect variation in musical/rhythmic expertise (e.g., Hove, Keller, &

Krumhansl, 2007).

In addition to testing the prediction above specifically, we have three questions of

interest that ‘tap’ into ongoing scientific endeavours. First, we will examine to what extent

microtiming influences the experience of groove and quality of sensorimotor synchronisation

(Danielsen et al., 2015; Davies et al., 2013; Kilchenmann & Senn, 2015). Second, we aim to

explore possible systematic relations between effort and groove experience, moderated by

rhythmic structural complexities and features. Third, by comparing professional musicians

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with non-musicians on the cognitive processing of polyrhythm and microtiming, the study

contributes to documenting effects of music exposure, music-related activity and music

training on brain plasticity, as manifested in behaviour, cognition, brain structure and

function (Aheadi, Dixon, & Glover, 2010; Angulo-Perkins et al., 2014; Gaser & Schlaug,

2003; Hansen, Wallentin, & Vuust, 2012; Hove et al., 2007).

The present study involves two experiments that were designed and carried out by the

author and supervisors. Both experiments feature a paradigm where music clips were

presented auditorily to demographically-matched groups of musicians and non-musicians.

Infrared eye-tracking equipment recorded participants’ pupil sizes during music listening. In

addition, in some conditions, the participants tapped the perceived beat on a key of the

computer keyboard and gave subjective ratings after the trials.

Following this brief introduction, we will continue by defining and elaborating on

concepts and terms necessary for a thorough discussion of the topics at hand. We look at

‘enculturation’ and active musical experience as premises for enabling and refining music

perception and prediction in general. Afterwards, we will exclusively focus on musical

rhythm and its relevant concepts. Theoretical frameworks on rhythm, beat and meter

perception are presented, before we examine research and concepts related to movement to

music, groove and the rhythmical features of polyrhythm and microtiming. The present study,

its goals and purpose, as well as the specific hypotheses, will be clarified in more detail,

followed by presentation of methods and results. Finally, we will discuss our results in light

of our research questions, existing theory and empirical data.

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2 Theoretical foundation 2.1 Music perception is enabled by enculturation

and refined by musical training ‘Engaging with music is the brain’s equivalent to a full body workout’. The words are taken

from a recent TED-education movie (Collins, 2014), summing up cognitive neuroscience

research of music from the last decades. Indeed, music listening constitutes the processing of

a continuous stream of complex auditory signals that apparently activates ‘the whole’ brain,

in particular the auditory association areas in the temporal lobe, auditory working memory

areas in the frontal lobe, and the emotional centres within the limbic system (Peretz &

Zatorre, 2005). Music performance is an even more demanding endeavour, termed one of the

most cognitively challenging tasks the brain can undertake – a complex and precise interplay

between auditory, somatosensory, motor, premotor and prefrontal areas (Klein, Liem,

Hänggi, Elmer, & Jäncke, 2016; Zatorre, Chen, & Penhune, 2007). The latter is also

considered a typically studied domain of expertise at both the cognitive and behavioural

levels (besides such activities as chess-playing, reading and sports; Ericsson, 1996).

The framework for understanding incoming local auditory events as meaningful musical

units develops from the very beginning of our life, as we (perhaps in concert with genetic

pre-dispositions) gain basic music perception skills from passive culture-specific music

exposure (enculturation; Hannon & Trainor, 2007). Thousands of hours of attentive but also

inattentive, incidental musical experience leaves us with an arsenal of tacit knowledge,

schemas and – importantly – predictions, concerning how our culture’s music normally

unfolds over time. For example, which tone combinations are most probable to occur, how

chord changes are tending to proceed, how rhythmic patterns are usually built up, and so on

(Huron, 2006; Pearce, Ruiz, Kapasi, Wiggins, & Bhattacharya, 2010). Exposure to these

statistical regularities of music ‘enculturates’ us into the music’s ‘grammar’, a process similar

to acquiring our mother tongue on the basis of being passively exposed to language (Sloboda,

1985). In addition, the research literature has extensively documented how active music

engagement changes the way music is perceived. It is well established that musical training

enhances domain-specific music perception, with evidence from behavioural studies (Hove et

al., 2007; Rammsayer & Altenmüller, 2006; Tervaniemi, Just, Koelsch, Widmann, &

Schröger, 2005) and brain imaging (Alluri et al., 2017; Fujioka, Trainor, Ross, Kakigi, &

Pantev, 2005; Vuust et al., 2005). Positive effects of musical training when dealing with

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auditory, non-musical tasks have also been reported (Aleman, Nieuwenstein, Böcker, & de

Haan, 2000; Martens, Wierda, Dun, De Vries, & Smid, 2015). In fact, musicians’ brains do

look different in connectivity and local cortical size than non-musicians’ brains (Gaser &

Schlaug, 2003), but remarkably, changes in neural connectivity in novices has been detected

already after a few months of attending regular drumming lessons (Amad et al., 2017).

Finally, music listening alone may have an enhanced effect on non-musical activities, like

spatial tasks (Aheadi et al., 2010), keeping perceptual awareness over time (Olivers &

Nieuwenhuis, 2005), and social cooperation (Anshel & Kipper, 1988).

2.2 Perception of rhythm, beat and meter Like the perception of all complex stimuli in our environment, perceiving music involves a

delicate interaction between bottom-up and top-down processes in our brains. Moving on to

the field of rhythm perception, the bottom-up/top-down interplay has been given considerable

research attention, especially between rhythm and meter. To understand the distinction

between the two, we observe that music is a temporal phenomenon and rhythm is often

defined as patterns of distinct time duration (‘inter-onset-intervals’) that are actually present

in sounding music (London, 2012). Its perception is assumed to be aided by innate capacities

like general gestalt principles of grouping, for example proximity, similarity and continuity

(Clarke, 1999; Jones, 2016). An organizing temporal mental framework in rhythm perception

is the one of sensed meter, which is a perceptual and cognitive phenomenon (Jones, 2016;

London, 2012; Palmer & Krumhansl, 1990). Sensitivity to meter is evident when we

spontaneously (and pre-attentively; Damsma & van Rijn, 2017) stomp – usually isochronous

– beats along with music, and forms our experience of timing regularities, including periodic

shifting between strong and weak beats. More extensively defined, meter may be understood

as a rhythmic virtual reference structure or point of departure, providing listeners and

performers with, in principle, any possible scaffolding aspect to make sense of the rhythmic

structure (Danielsen, 2018; Danielsen et al., 2015). This includes idiosyncratic patterns of

individual top-down matrices that may also involve non-isochronous temporal relations. To

clear up the terminology, we will in the following understand meter as an internal framework

when attending to rhythm, consisting of hierarchically grouped beats with related (also non-

isochronous) subdivisions. A related concept to meter perception, beat perception (Honing,

2012; also termed beat induction), addresses the inferring of beats only – a detection of a

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regular pulse in an auditory signal (Winkler, Háden, Ladinig, Sziller, & Honing, 2009),

without subdivisions or hierarchical groupings.

An experience of beat and meter is fundamental to our capability to synchronise to

musical rhythm. Different metrical frameworks of the same rhythmic material represent

different perceptual experiences (Iversen, Repp, & Patel, 2009) and an absence of a sense of

meter contributes to the feeling of ‘being lost’, i.e., not being able to grasp what is

rhythmically going on. Importantly, the internal metric framework does not need to actually

be played to be comprehended by the listener; it may simply be evoked by cues within the

rhythmic texture (Honing, 2012).

The perception of beats has been linked in neuroscience to activations within basal

ganglia and cerebellum, primary, premotor and supplementary motor areas (Grahn & Brett,

2007), while the processing of metric aspects is especially associated with activity in

language areas of the inferior frontal gyrus (BA47; Vuust, Wallentin, Mouridsen, Østergaard,

& Roepstorff, 2011). There have also been experimental attempts to reveal neural substrates

related to the processing of syntactical complexity (syntax here refers to the ways in which

the notes of the pattern are rhythmically combined) in rhythmic structure (i.e., rhythmic

patterns; more related to bottom-up than to top-down processes). In a recent study aimed at

investigating brain response to increase in syntactical complexity in a drum groove, brain

areas were found to include the left cerebellum, the right inferior frontal gyrus (comprising

BA47) and superior temporal gyri on both sides (Danielsen, Otnæss, Jensen, Williams, &

Østberg, 2014). Interestingly, it has been suggested that syntactical processing in language

and music partly shares neural resources (Patel, 2003).

2.2.1 Meter perception I: Metric Encoding Theory and Dynamic

Attending Theory Several cognitive theories have accounted for our ability to infer and experience meter from

sounding rhythm. The Metric Encoding Theory (Povel & Essens, 1985) suggests that

listeners sense of meter arises from cues in the actual pattern of sounds, often regular accents

in the subdivisions. A series of ‘mental beats’ are induced – what the authors call an ‘internal

clock’ (Abernethy, 1988; Povel & Essens, 1985). This inner series of beats will continue to

be endorsed, until the sounding music’s accents contain too many counter-evidence events,

i.e., too little clock-fit. At this threshold, the listener’s sense of meter is given up in favour of

another reference structure, or possibly a feeling of confusion or just ‘float by’. The Metric

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Encoding Theory has been criticized on several points: for its ‘rigidity’ in describing inner

beats as a stable clock with fixed lengths between beats (not adjusting to timing variations),

in taking into consideration too little the use of the body in the process, and in not positing

multiple time level relationships as aids in meter perception (for an extensive review of

critique, see Jones, 2016).

The Dynamic Attending Theory (DAT; Jones, 2016; Jones & Boltz, 1989) addresses

all of the above aspects. With support from neuroscientific evidence, DAT claims that as

external rhythmic events are paid attention to, neural population oscillations are set into

action, defined as rhythmic or repetitive activity (Fujioka, Trainor, Large, & Ross, 2009;

Large, Herrera, & Velasco, 2015; Lehmann, Arias, & Schönwiesner, 2016; Nozaradan,

Peretz, Missal, & Mouraux, 2011). These neural oscillations are closely connected with overt

body movement, as will be addressed later, and correspond to what in DAT is termed the

attending rhythm. Specifically, the oscillations represent multiple time levels of what is heard

– rhythm – or imagined through a metric framework (Iversen, Repp, & Patel, 2005).

Furthermore, being self-persistent, the oscillations ‘expect’ stability. Their cyclic nature

entails anticipations concerning the placement of the next pulse beat and hence, cues to the

temporal allocation of attentional energy (Jones & Boltz, 1989). Such anticipations help us

direct our attention adaptively, so that the efficiency of the processing of the incoming

stimulus is increased (Greenberg & Larkin, 1968). DAT proposes that high synchronisation

of the oscillations with the external sound source sharpens the attentional focus, giving a

higher degree of expectancy violations when perturbations occur. Expectancy violations, e.g.,

small timing asynchronies, are continuously taken into account, widening the attentional

focus for each perceived beat in order to encompass deviations or onsets, or adjusting the

oscillation phases (Danielsen et al., 2015). Importantly, although not stated explicitly by the

theorists, we may assume that such adjustments demand attentional effort.

The hypothesis of metric binding extends DAT (Jones, 2016). When different

oscillations are internally co-activated over time, association links are formed, binding the

oscillations together. This contributes to the listener’s internalizing of, and familiarity with

different meter and rhythm categories. The persistence of these ‘binded’ oscillations is

dependent on their phase relationship fit and the listener’s ability to predict the following

rhythmic events. Some oscillation phase ratios, for instance 2:1, are more stable than 4:3 (a

polyrhythm, see section 2.4.1) and require less time for the nesting to take place. Extensive

musical experience helps listeners both mentally activate complex oscillation ratios

(Stupacher, Wood, & Witte, 2017), and flexibly alternating their attention between

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oscillations at different time levels (Palmer & Krumhansl, 1990) – also when oscillations are

not ‘supported’ in the external music. It may be assumed that activating and maintaining

complex oscillation ratios generally demands attentional effort on the part of the listener. As

its name implies, the Dynamic Attending Theory is indeed a dynamic one, emphasizing the

top-down/bottom-up interaction of perception of rhythm, with continuous adapting to the

nature of the external input.

2.2.2 Meter perception II: Predictive Coding Theory As it was suggested both in the sections on enculturation and DAT, anticipation is an

important part of music perception. In fact, Huron (2006), building on Leonard B. Meyer’s

classic theory (1956), places the anticipations, fulfilments and violations of these at the very

heart of our aesthetic experience and emotional response to music. The broadly scoped

Predictive Coding Theory (PC; originally proposed by Friston, 2005) takes the notion of

prediction further in explaining rhythm and meter perception (Vuust & Witek, 2014). The

main assumption is that the brain generally operates with the ultimate goal of ‘predicting the

future’. Taken into a rhythm context, incoming (rhythmic) stimuli are continuously compared

or matched against a mental model (meter) formed from enculturation and formal musical

training. Mental models give rise to anticipations of what will happen next in the meeting of

sensory stimuli and contextual information. Different models might compete in making sense

of the incoming rhythms, and the internal model that is favoured tends to be that which best

anticipates incoming data. In other words, PC claims that meter is the internal cognitive

model that ‘minimizes the error’ between the brain’s predictions and the incoming rhythmic

surface structure. As in DAT, mental models are dynamic and open for in-the-moment

updates with the incoming musical information, and the prediction error and following

adjustments and updates are assumed to demand effort.

2.2.3 Development and neural basis of beat and meter perception With evidence from brain activity measures, Winkler et al., (2009) showed that newborn

infants are able to perceive the beat from incoming rhythm. Similarly, experiments

employing habituation/dishabituation-paradigms that associate auditory sequences with

visual fixation times have demonstrated that infants of only two months of age discriminate

between series of isochronous beats with small tempo differences (Baruch & Drake, 1997)

and by 7 months infer metrical frameworks from rhythmic patterns (Hannon & Johnson,

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2005). On the basis of findings like these, it has been argued that innate predisposition alone

can account for the perception of beats (Honing, 2012). However, it is also likely that sources

of enculturation may provide substantial contributions; unquestionably, when it comes to

meter perception skills, experience seems to be a crucial component for development and

shaping. Important enculturation factors are pre- and post-natal exposure (imprinting) to

basic biological rhythms such as breathing (0.1-1 Hz), walking (1-3 Hz) or the rhythm of

speech (3-10 Hz) in caregivers and others (Hannon & Trainor, 2007; Iyer, 2002), as well as

culture-specific aspects. Effects of enculturation have been found in empirical investigations.

For example, according to Hannon & Trehub (2005), by six months of age, infants respond

similarly to disruptions of rhythms from different cultures (Western versus Balkan music),

but by one year of age they respond more readily according to their own culture’s rhythmic

systems. Furthermore, Phillips-Silver & Trainor (2005) demonstrated the influence of

multimodal caregiving on meter perception. Infants that over a period of time had been

rocked by their mothers, but only according to every second or every third beat in a non-

accentuated pulse train, later showed a preference for accents that corresponded to their

‘rocking’ rhythm pattern (every second or third beat).

If enculturation shapes the perception of meter, does extensive musical training enhance

it? Some results have suggested that elementary (Western) meter perception is independent of

musical sophistication (Bouwer, Van Zuijen, & Honing, 2014; Damsma & van Rijn, 2017),

while others suggest that musicianship and expertise fundamentally develop metrical

frameworks enhancing perception of rhythm (Chen et al., 2008; Drake, Penel, & Bigand,

2000; Geiser, Sandmann, Jäncke, & Meyer, 2010; Matthews, Thibodeau, Gunther, &

Penhune, 2016; Stupacher et al., 2013, 2017; Vuust et al., 2005). When it comes to more

complex rhythmic structure (after Western standards), musicians have indeed been found to

be more sensitive than non-musicians in perceiving and/or superior in synchronising to

complex meters (Snyder, Hannon, Large, & Christiansen, 2006), musical syncopations

(Ladinig et al., 2009), polyrhythmic musical texture (Jones, Jagacinski, Floyd, & Klapp,

1995) and onset (microtiming) asynchronies (Hove et al., 2007).

2.3 Synchronising to musical rhythm Music and movement are strongly interrelated dimensions and music typically co-exists with

dance (Brown, 2004). As mentioned in the introduction, sensorimotor synchronisation is the

coordination of rhythmic movement with an external rhythm (Repp & Su, 2013), observed

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even in toddlers (Kirschner & Tomasello, 2009), and mostly considered an ability of humans

(Wallin & Merker, 2001; but for evidence in other animals see: Fitch, 2013; Patel, Iversen,

Bregman, & Schulz, 2009). The concept of entrainment is built on DAT and provides a

theoretical account, as it refers to the process where two different oscillations are

synchronised – often the internal attending rhythm (i.e., neural oscillations) with an external

source (i.e., sounding music; Jones, 2016). Some authors theoretically separate mental

models and body movement, describing how bodies are coordinated with the external sounds

but synchronised with the internal model that predicts the beat and meter of the music (Repp

& Su, 2013). An embodied perspective on cognition, on the other hand, places body activity

at the heart of rhythm perception (Iyer, 2002). According to this perspective, perception and

cognition are assumed to be shaped by body activity. There are even (philosophical)

proponents, claiming that rhythm is experienced and grasped actively, non-analytically and

non-cognitively, within the body itself (Roholt, 2014). As dance and movement to music

involves the whole body, there is a growing scientific enthusiasm in quantifying body

movement trajectories with, for example, motion sensors (Jensenius, 2009). However, most

research on sensorimotor synchronisation has so far been done in the form of finger-tapping

paradigms (see Repp & Su, 2013 for a review of recent findings), where the participant is

typically asked to intentionally tap a finger in synchrony with isochronous beats.

2.4 Groove The concept of groove captures the three aspects of sounding rhythmic properties,

embodiment and pleasure (Witek, 2017). First, grooves are musical patterns that have a

certain rhythmic, symmetrical, continuously repeating and danceable quality to them

(Pressing, 2002). Groove-based genres include the ‘Black Atlantic rhythms’ (jazz, soul,

reggae, hip-hop, funk; Pressing, 2002) and contemporary computer-programmed styles (e.g.

electronic dance music, EDM; Butler, 2006). Second, groove can also be seen as a

psychological construct, being a subjective, affective and sensorimotor response to groove-

based music. One definition, ‘wanting to move some part of the body in relation to some

aspect of the sound pattern’ (Madison, 2006, p. 201), treats groove purely as a sensorimotor

phenomenon, whereas another, ‘the groove is that aspect of the music that induces a pleasant

sense of wanting to move along with the music’ (Janata et al., 2012, p. 56), adds in the third

aspect: the affective component. Related to the latter, Danielsen (2006) argues that presence

and pleasure are important characteristics of a groove experience, while Roholt (2014) on his

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side, discusses how groove may evoke feelings of tension and relief, depending on how

rhythms are experienced in the body.

Groove-based music is intended to make us entrain to the beat, especially activating

the locomotion (‘walking’) part of the sensorimotor system (Iyer, 2002). A strong link has

been found between a feeling of being ‘in the groove’ and the wish to synchronise body

movement to music (although movement constraints do not necessarily affect groove ratings;

Janata et al., 2012). It has similarly been reported that people move more (head and feet) to

high-groove tunes than low-groove tunes (Janata et al., 2012). However, the relation may not

be one-to-one, since Kilchenmann & Senn (2015) found that expert musicians moved more to

a rhythm excerpt that they rated less groovy. The authors suggested that musicians in this

situation used their body primarily to compensate for a fuzzy rhythmic situation and not to

respond to a groove experience. Despite the fact that the groove term is especially used in

relation to Afro-American and derived music heritage, a proposal has been made that

experience of groove is nevertheless a cross-cultural phenomenon (Madison, 2006). There

may, however, be individual and culture-specific differences in being able to perceive

particular grooves in particular musical styles (Roholt, 2014), again emphasising an

important role of culture-specific enculturation.

2.4.1 Two structural properties of groove-based music Which are the metrical, rhythmic and microrhythmic features of a groove that are actually

making it groove? The laborious work of unlocking the structural secrets behind groovy

music has begun to pay off. It turns out that some elements are related to structure that

affords successful entrainment, including temporal information that increases rhythmic

predictability, such as a ‘locomotion-friendly’ tempo (Janata et al., 2012), repetition

(Danielsen, 2006), structural ‘low-level’ features like event density (‘the density of sound

events between beats generally’; Madison et al., 2011, p. 1579), and beat salience ('the degree

of repetitive rhythmical patterning around comfortable movement rate'; Madison et al., 2011,

p. 1581). At the same time, however, it seems essential that certain forms of complexity that

challenge the listener’s perception of, and create tension in relation to, the sensed meter, are

also some of the ingredients. Regarding this, researchers have investigated and discussed, for

example, the role of syncopation density (‘changes in rhythmic emphasis from metrically

strong to metrically weak beats’; Witek et al., 2014, p. 2: Sioros, Miron, Davies, Gouyon, &

Madison, 2014), polyrhythm (Danielsen, 2006; Vuust et al., 2014) and microtiming

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(Butterfield, 2010; Danielsen et al., 2015; Iyer, 2002). The latter two – polyrhythm and

microtiming – are especially dealt with in the subsequent sections, as they are central topics

of the present study.

Polyrhythm

Polyrhythm refers to two (or more) beat levels, played by one or more performers, elapsing

simultaneously within a single rhythmic texture, for example 3:2 (‘3 against 2’), 2:3, 4:3, 3:4,

5:3 and 5:4. Of several types of polyrhythm, one is cross-rhythmic counter-rhythms

(Danielsen, 2016, p. 62; from now on termed 'cross-rhythms') where two (or more)

competing pulses appear in a single musical context; when experienced within a metric

framework, one is perceived as the main beat level and the other as the counter-rhythm.

Cross-rhythms are extremely common in groove-based music. In West African traditions for

example, continuously competing reference structures appear as a rule. The characteristic

12/8 rhythms can be understood in cycles of 3, 4, 6 or 8 (e.g., in Salif Keita’s 1987-tune

Wamba). In contemporary EDM, integrated two-bar 4:3 cross-rhythmic patterns are typical,

for example in Till The World Ends (2011) by Britney Spears or Where Are Ü Now (2015) by

Skrillex. Similarly occurring multiple double- and half-time-feels are also common, as in

Destiny’s Child’s Nasty Girl (2001; analysed by Danielsen, 2015). Also in jazz, cross-

rhythms are an essential part of the musical language. Vuust & Roepstorff (2008) gave an

extensive analysis of polyrhythm in Miles Davis’ 1960s quintet.

There are clear analogies in perceptual

effects between auditory cross-rhythms and

some forms of so-called ‘illusory’ perception,

for example, bistable visual percepts (Jastrow,

1899). The picture shown in Figure 2 may be

seen either as rabbit or a duck, giving rise to two

completely different, and mutually exclusive,

gestalts. Similarly, cross-rhythms are bistable

and ambiguous auditory percepts, that will be

experienced differently depending on which

internal metric framework is endorsed; that is, which beat level is perceived as the main level

and the counter-rhythm, respectively (Vuust & Witek, 2014). Although it is possible to hear

both competing pulses at the same time, or alternate between them, simple polyrhythms are

often heard as a merged rhythmic texture, not as two separate streams (Deutsch, 1983; Kurtz

Figure 2. A bistable percept. From Jastrow (1899).

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& Lee, 2003). Keeping and sticking to a given metrical model is possible both when the

acoustic salience is on the counter-rhythm, and even when the meter is not acoustically

actualized at all. The latter, however, is considered a cognitively demanding task, since it

forces a beat level that is not present in the music, to be internally maintained.

Microtiming

The tones, chords and hits played by musicians are typically not in full temporal synchrony

(even if musical training greatly helps; Gérard & Rosenfeld, 1995). Asynchronies emerging

from limitations in perceptual, cognitive and motor faculties have been characterized as

‘motor noise’ and ‘discrepancies’ (Rasch, 1988). However, certain forms of microtiming,

subtle timing asynchronies in music, are also systematically and intentionally (although not

always consciously) applied for expressive purposes (Bengtsson, Gabrielsson, & Thorsen,

1969; Clarke, 1989; Collier & Collier, 1996; Danielsen et al., 2015; Iyer, 2002; Keil, 1987).

The musician or ensemble may play rhythmically ‘outside’ a presumed norm (the norm most

commonly constitutes an isochronous beat series/metric grid), there may occur subtle tempo

changes or there may be asynchronies between the onsets of the player’s or players’ tones.

For example, in some pianists’ solo performances (Palmer, 1996) and string trios (Rasch,

1988), the melody line is often played 5-50 ms ahead of the rest of the parts. Jazz players can

be distinguished by their personal rhythmic signature – ‘feel’ – that gives to each of them a

coherent and recognizable delivery of notes and phrases (Iyer, 2002; Pressing, 2002). In jazz

drumming, microtiming of cymbal swing ratios (short-long eighth-note patterns) has been

extensively studied (Friberg & Sundström, 2002). Consistent timing asynchronies have been

discovered between piano, drums and bass in typical swing accompaniments (Collier &

Collier, 1996). Also, ‘trade-off’-effects in terms of the relative lengths of the beats in 4/4 bars

(hierarchical groupings of four and four beats) are common when jazz bands perform swing

pieces (Ross, 1989, from Collier & Collier, 1996); some beats (1st and 3rd) being consistently

shorter and some beats (2nd and 4th) consistently longer. In R&B music, artists like D’Angelo

have taken microtiming to an extreme level, including systematic timing asynchronies

between the drums and electric bass of up to 90 ms (Danielsen, 2010a; Danielsen et al.,

2015). Also, in contemporary groove based computer-made music, microtiming has become

an integral part of the rhythmic fabric, where timing nuances can be programmed in Digital

Audio Workstations, without expert instrumentalists having to play them (Danielsen, 2010b,

2015). Interestingly, in some traditions, for example drumming music from Mali (Polak,

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2010) and Norwegian fiddle music (Haugen, 2015), a temporal hierarchic reference structure

is lacking and microtiming ‘deviations’ constitute the actual norm.

Remarkably, despite being a seemingly essential component of musical rhythm,

microtiming is often not consciously perceived (Hove et al., 2007). Still, there is reason to

believe that notes containing subtle timing discrepancies will stand out from notes placed on

the rhythmic grid, tune our attention towards them, and our perceptual system will treat them

as something worth a closer analysis (Iyer, 2002). Musically, this might help reaching a

greater perceptual salience for some structural aspects of the music, like melodic material or

even single tones (Iyer, 2002; Palmer, 1996). The Participatory Discrepancy Theory (Keil,

1987; Keil & Feld, 1994) suggests that subtle timing asynchronies are also one of the active

components contributing to the experience of groove. Keil (1987, p. 277) writes: ‘It is the

little discrepancies between hands and feet within a jazz drummer’s beat, between bass and

drums, between rhythm section and soloist, that create the swing [e.g., groove] and invite us

to participate’. This is in line with for example jazz musicians’ first-hand experience (see

Collier & Collier, 1996), often discussing rhythmic nuances like ‘pushing’, ‘pulling’, or

playing ‘laid back’ or ‘on the beat’ when playing groove-based music. Similar to Keil, Witek

(2017) suggests that rhythmic complexity (mainly discussing musical syncopations) requires

a certain degree of active participation on the part of the listener to bodily ‘fulfil’ the groove.

The same can perhaps be said for microtiming (and also polyrhythm), as it sets up patterns of

tension and equilibrium toward listeners’ anticipations of a metrical pulse (Roholt, 2014).

This may have behavioural implications, as Danielsen and colleagues (2015) found a

qualitative change in bodily entrainment with a rhythm when a microtiming pattern was

introduced: bodily synchronisation was drawn towards the added asynchrony-onset.

Moreover, they saw a more pronounced entrainment to the half note level of the rhythm,

probably because more complex microtiming obscured the exact location of beats at several

quarter note positions. Both were assumed to be signs of the metric adjustment following the

challenging of the listeners’ fine-grained temporal expectations. Importantly, though, despite

extensive theorizing and anecdotal evidence pointing to the causal link between the structural

element of microtiming and groove experience, experimental studies investigating this notion

have revealed inconsistent results (Butterfield, 2010; Davies et al., 2013; Frühauf, Kopiez, &

Platz, 2013; Kilchenmann & Senn, 2015; Senn, Kilchenmann, von Georgi, & Bullerjahn,

2016).

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2.5 Pupillometry: a measure of cognitive processing Until now, we gave a short introduction on enculturation and musical training’s effects on

music perception in general, before we narrowed the focus to musical rhythm. We looked at

rhythm, beat and meter perception as well as body movement to music, the concept of groove

and two structural features of groove-based music, namely polyrhythm and microtiming. As

was discussed, both DAT and PC imply some degree of mental effort when perceptually

processing (or playing) rhythm in relation to a corresponding meter, especially when the

metric model is challenged and there is need for internal adjustments. It was also discussed

how such a pressure of metric models, generated by certain rhythmic complexities, in some

instances seem to promote the groove experience. Finally, many findings have pointed to the

more sensitive and refined rhythm and meter perception skills of musicians. Such skills may

possibly influence the mental effort recruited when perceiving rhythm and rhythmic

complexity in relation to metric frameworks. Therefore, it may be very relevant to investigate

‘musical effort’ in musicians and non-musicians, related to the processing of polyrhythmic

and microtiming events.

In his classic book Attention and Effort, Kahneman (1973) first proposed the

psychological concept of mental effort, a dimensional construct that he claimed was most

precisely measured by monitoring diameter changes of the eye pupil – a method typically

called pupillometry (Laeng et al., 2012; see also Kahneman, 2011, chapter 2). At the time

Kahneman’s book was released, psychological experiments had already been using

pupillometry for about two decades. The very first studies applied the methodology to

measure people’s pupillary responses in relation to affective arousal, mainly from pictures

(Hess & Polt, 1960). Later it became clear that changes in pupillary size could also reflect the

‘intensive’ side of attention, the capacity facet rather than the selection facet (Hess & Polt,

1964; Laeng et al., 2012), namely mental effort in Kahneman’s terms of allocation of

cognitive resources to a current mental task or behavior. Implicit in this view is that brain

processing resources are continuously allocated via top-down attentional processes towards

task-relevant stimuli in order to meet current demands. Mental effort refers to variation in

those processing demands (e.g., Alnæs et al., 2014). In other words, increased pupil size

reflects an increase in mental (cognitive) workload (Lean & Shan, 2012). Importantly, a same

task may result in different levels of mental effort, on the grounds of individual differences in

ability, cognition, and temporary psychological and physiological state (Lean & Shan, 2012).

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In recent years, it has become clear that the brain regulates the level of processing

resources via changes in arousal in various systems of the brainstem. The locus coeruleus

(LC) is a key arousal system that appears specifically involved in cognitive ‘arousal’ more

than vigilance or waking state per se. The LC is also the noradrenergic (NE) ‘hub’ of the

brain and it is located in the pons on both sides. Recent research confirms that the LC-NE

system regulates noradrenergic neuromodulation and it concomitantly causes pupillary

changes in relation to these cognitive operations (Joshi, Li, Kalwani, & Gold, 2016).

Experiments have revealed high correlations between pupil response and neuronal activity in

the LC (Rajkowski, Majczynski, Clayton, & Aston-Jones, 2004). Pupil dilation may reach up

to 20% from baseline levels as a result of cognitive processing alone, and some strong

feelings like pain and orgasm may dilate the pupil even more. However, variations from

illumination can account for about 120% dilation change (Laeng et al., 2012). Therefore,

pupillometry must be conducted in luminance-controlled conditions and with specialized

equipment. Most cognitive-related changes of the pupil are difficult to see with the naked

eye, and they may be ‘swamped’ by luminance change in daily contexts.

The pupil response has been found to index external events with high temporal

resolution (Damsma & van Rijn, 2017; Laeng, Eidet, Sulutvedt, & Panksepp, 2016).

Furthermore, as Laeng and colleagues (2012) point out, cognitive processing does not need to

be conscious to be reflected in pupil size. For example, amnesics (Laeng et al., 2007) and

blindsight patients (Tamietto et al., 2009) respond with variations in pupillary diameters even

when lacking conscious awareness of exposure to significant stimuli. It seems also difficult to

control pupil size voluntarily (Laeng et al., 2012), which makes the pupil a particular reliable

(honest) signal of the underlying processing.

Measures of pupillary response have been shown to be an indicator of several

attentive and pre-attentive cognitive mechanisms, including interest and affective processing

(Laeng et al., 2016; Partala & Surakka, 2003), processing of language (Schluroff, 1982),

surprise and violation of expectation (Damsma & van Rijn, 2017; Friedman, Hakerem,

Sutton, & Fleiss, 1973), cognitive conflict/interference (Laeng, Ørbo, Holmlund, & Miozzo,

2011), working memory load (Granholm, Asarnow, Sarkin, & Dykes, 1996), and

perceptual/attentional shifts (Einhäuser, Stout, Koch, & Carter, 2008).

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2.5.1 Mental effort in the study of rhythm and meter perception A common approach in the study of brain activity related to rhythm and meter perception has

been the mismatch-negativity (MMN) component of event related potentials (ERPs) in

electroencephalography (EEG; Näätänen et al., 2007), where magnitude and latency of early

EEG-signals index a mismatch between expectation and what is heard (e.g., Honing, 2012;

Vuust et al., 2005). The methodology has been used for example to test specifically PC

theory, where typically single time-locked rhythmic ‘oddballs’, that is, rhythmic deviants or

omission of one or more beats, have been linked to an electrophysiological response. The

response has been interpreted as degree of expectation violation between rhythm and

underlying meter. Similar to the MMN component, time-locked pupil dilations have also

been employed to signal metric violation/‘surprise’ effects to rhythmical deviants (Damsma

& van Rijn, 2017). In some studies, stronger responses to single rhythmic deviants in

musicians than non-musicians have been observed, which have been accounted for by the

enhanced sensitivity of musicians to musical incongruity, caused by stronger metric models,

and seemingly supporting PC (Brattico et al., 2008; van Zuijen, Sussman, Winkler, Näätänen,

& Tervaniemi, 2005; Vuust, Ostergaard, Pallesen, Bailey, & Roepstorff, 2009).

As for the present study, an alternative approach to EEG was adopted, namely monitoring

time-averaged pupillary diameters while participants listened to continuously running

rhythms with varying degrees of structural complexity, but without time-locked, single

deviants. We decided to employ this approach to provide a measure of attentional demands

(e.g., mental effort) as participants adjusted their internal metrical model over a period of

time in accordance with the incoming rhythmic texture in order to reduce mismatch error, as

DAT and PC specifically propose. An advantage is that the present paradigm encompasses

both the initial response (e.g., prediction error) and the following adjustment of the metric

framework to ‘fit’ the incoming rhythm, and may therefore capture a different process than

for example the MMN-studies above. Individual and group (musicians vs. non-musicians)

differences in effort, as revealed by the pupils, may be interpreted as differences in perceptual

ability, revealing that differently experienced brains indeed process and analyze musical

information differently.

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3 The present study Two experiments were designed and carried out to explore professional jazz musicians’ and

non-musicians’ processing of musical polyrhythm and microtiming in groove contexts. Our

dependent variables consisted of three complementary data sources (Jack & Roepstorff,

2002), covering several aspects of pre-attentive and conscious processing: 1) Levels of

mental effort, as measured by group-averaged pupil diameter change; 2) Quality of

sensorimotor synchronisation (tapping accuracy), operationalized as standard deviation of

tapping offset from rhythmic reference points; and 3) Subjective ratings (in The Microtiming

Experiment only) of groove (‘wanting to move the body to the music’) and ‘musical well-

formedness’. All musical stimuli were presented in a passive (‘listening only’) condition and

an active (‘synchronising with the beat’) tapping condition.

For The Polyrhythm Experiment (Experiment 1), we borrowed from the paradigm

used by Vuust and colleagues (2006) two 30-second excerpts from a single ‘real’ musical

track, namely Sting’s tune Lazarus Heart (1987). Specifically, we wanted to investigate how

cross-rhythmic events (4:3) influenced pupillary change and tapping accuracy compared to a

comparable but non-polyrhythmic excerpt. In The Microtiming Experiment (Experiment 2),

we exposed participants to 30-second long original double bass and drum grooves of varying

structural complexity by increasing the number of musical syncopations and note onsets per

bar into three complexity levels. Within two of these three levels we compared the same

grooves with and without hi-hat eighth notes, to directly measure a ‘Timekeeping’ effect,

being a type of ‘event density supporting metric reference’. Finally, for all groove excerpts,

we systematically changed the asynchrony magnitude between the double bass and the drums

into five distinct microtiming conditions. We were mainly interested in microtiming, but also

how other rhythmic features (syncopation/note onsets per bar and timekeeping) were related

to effort, sensorimotor synchronisation and subjective ratings.

3.1 Hypotheses and predictions Our general hypotheses concerning effort and tapping accuracy were partly generated by the

two influential theories Dynamic Attending Theory (DAT) and Predictive Coding Theory

(PC). However, the present study should be seen primarily as exploratory, since previous

experiments on the same topics are lacking. Also, predictions derived from the above theories

seem somewhat unclear with respect to several aspects of the present stimuli material and,

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most importantly, the two theories generate few obviously competing predictions concerning

effort in relation to rhythm and meter perception. However, it is reasonable to argue that

according to both PC and DAT, the degree of correspondence between metric framework and

continuously incoming rhythm is negatively related to both processing intensity within the

perceptual system (mental effort) and sensorimotor synchronisation accuracy. Hence, when

specific rhythmic instances challenge listeners’ metric frameworks, this will decrease the

meter/rhythm correspondence, thus increase prediction error and concomitantly the need for

adjustment. This would happen in order to fit the metric framework to the incoming rhythm,

as both theories propose, compared to when meter is not challenged by such rhythmic

features. Increased prediction error and internal adjustment demands should together yield

increased mental effort and decreased sensorimotor synchronisation accuracy.

Polyrhythm and microtiming are rhythmic features that are assumed to challenge

metric frameworks, as they both deviate from a ‘normal’ on-the-grid rhythmic texture, built

around a single and unambiguous pulse. In Experiment 1, listeners will face a competing and

distracting counter-rhythmic pulse level on ‘top’ of an acoustically absent main beat level

(i.e., cross-rhythms) that disturbs their fundamental sense of beat and meter. Hence,

maintaining the original beat level is a cognitively demanding task. As for Experiment 2, the

task is more related to temporally placing the beat exactly than internally maintaining a beat

level. In other words, what represented the challenge when exposed to microtiming,

according to DAT and PC accounts, are the (micro)-adjustment of participants’ mental metric

framework/oscillation phases and/or wideness of attentional focus to encompass the onset

asynchronies. To summarise, we predicted that the cognitive processing of polyrhythm and

microtiming in groove contexts should increase pupil dilation and decrease tapping

accuracy, compared to when such rhythmic instances are not present.

In accordance with extensive empirical research documenting effects of musicianship,

it can also be claimed on the grounds of DAT and PC that musicians possess stronger, more

refined metric models than non-musicians. We expected this to be manifested by consistently

higher tapping accuracy in musicians than non-musicians. Our hypotheses are however less

clear in terms of possible pupillary effects of rhythmical expertise. On one side, stronger

metric models could result in higher perceptual sensitivity and hence more prediction error

when faced with rhythmic perturbations and complexities. On these grounds, one would

expect pupil dilation to be larger for musical experts. This is in line with the MMN-studies

that found stronger brain activation in musicians to single metric instances of rhythmic

incongruity. On the other hand, musicians could also be more familiar with and thus more

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tolerant to metric challenge and metrical ‘counter-evidence’ than non-musicians. Therefore,

they would perhaps adapt and adjust faster and more efficiently their internal metric models

to continuous and repeating musical input, using less effort over a period of time. The

prediction in terms of possible pupillary effects of greater tolerance to metric challenges is

therefore a larger pupil dilation in the non-musicians.

Since tapping involves motor response in addition to listening, we expected the active

condition to generate higher effort measures than the passive condition, which should be

expressed in greater pupil dilations in the former than in the latter. Finally, in line with

earlier research, we expected that the on-the grid or very moderate microtiming versions

would be rated highest on subjective measures of groove and musical well-formedness.

Nevertheless, we expected a group difference also on these ratings, as musicians’ stronger

metric models should be more adjustable to subtle onset asynchronies in timing (Hove et al.,

2007).

3.2 Methods 3.2.1 Participants

We recruited 32 volunteer, unpaid, participants (mean age = 30.06, range = 20-44). Sixteen

of the participants were professional jazz musicians and another 16 were non-

musicians/amateur musicians. In each group, there were 9 males (musicians: mean age =

32.1, range = 20-44; non-musicians: mean age 32.2, range = 20-44) and 7 females

(musicians: mean age = 27.4, range = 24-31; non-musicians: mean age = 27.3, range = 23-

31), giving a total of 18 males and 14 females. Each professional jazz musician was matched

with a non-musician of the same age (±2 years; so that the mean age of the two groups did

not differ), sex and completed level of education (in total, two participants had high-school

degrees, 24 had a Bachelor degree and 10 had a Master’s degree). All professional musicians

had their university degree in music and characterized themselves as jazz musicians, namely:

one trombonist, two sax players, three trumpeters, three pianists, two vocalists, two guitarists,

one bass player and two drummers. Among the non-musicians, five participants played as

amateurs the guitar and one played the flute. No other demographic data were recorded. All

participants had normal or corrected-to-normal (with contact lenses) eyesight and hearing

ability within the normal range, by self-report. Participants from both groups were recruited

by personal invitation from the author or word-of-mouth. All participants were able to

complete the experiment and signed a written informed consent before participation.

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Musical Ear Test (MET)

To verify objectively that the two groups of musicians and non-musicians differed in musical

expertise, in particular in relation to rhythm, a reliable and valid score of rhythmic

competence was applied. Specifically, all participants went through the Musical Ear Test

(MET; Wallentin, Nielsen, Friis-Olivarius, Vuust, & Vuust, 2010). MET has a melodic and a

rhythmic part (as for the present study, only the rhythmic part was used). The test consists of

52 pairs of rhythmic phrases, each phrase one bar long, consisting of 4-11 wood block beats

in tempo 100 bpm. The two phrases are presented immediately after one another, and the

participants’ task is to decide whether the pairs are identical or not; 26 identical and 26 non-

identical phrase pairs being included (see Figure 3).

Figure 3. Example from Musical Ear Test, rhythm part. From Wallentin et al., (2010).

Despite MET does not address competence of polyrhythm and microtiming

specifically, it measures a general musical/rhythmic competence through the proxy of

musical working memory. Thus, the test seems able to discriminate participants’ sensitivity

to auditory fine-scaled rhythms (Wallentin et al., 2010), which fits the goal of the present

study. MET has shown good psychometric attributes with practically no ceiling- or floor

effects. Moreover, it has been shown to successfully distinguish between groups of

professional musicians from amateur musicians and non-musicians (for further information,

see Wallentin et al., 2010).

A one-way analyses of variance (ANOVA) indicated a significant effect, F(1, 30) =

17.05, p < .001, of Musicianship (musicians vs. non-musicians) for the % correct responses

in the Musical Ear Test, rhythm part. This was due to the expected difference between the

musicians (M = 87.98; SD = 4.63) outperforming the non-musicians (M = 78.37; SD = 8.08)

on this test. For the musicians only, simple linear regressions were calculated to predict

MET-score based on their reported Hours of practice (daily on their instrument), and what

Age they began to play their instrument. Both analyses failed to reveal significant

relationships.

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3.2.2 Stimuli Experiment 1: The Polyrhythm Experiment

Experiment 1 consisted of two 30-second excerpts from the sax solo of Sting’s tune Lazarus

Heart (1987); one excerpt included polyrhythmic texture, the other did not and served as a

control stimulus. Stimuli were borrowed from Vuust et al. (2006); however, the stimuli’s

duration was shorter in the present study than in the original. Excerpt 1, ‘the polyrhythmic

condition’, contained three 4/4 bars of main meter (M), in tempo 120 bpm, followed by three

(main-meter) bars with a ‘competing’ counter-rhythmic beat level (C), 4:3 (the counter-

rhythm tempo being 160 bpm), without the main meter being explicitly musically referred to.

This sequence was repeated in order to give the pattern of M-C-M-C-M. In Excerpt 2, ‘the

plain condition’, C was excluded, so M was repeated five times (M-M-M-M-M). The main

meter sequence M and cross-rhythmic sequence C were of almost identical loudness, tempo

and length (six seconds), making the excerpts ideal for studying the effect of a polyrhythmic

texture in a musical, ecological setting. Figure A1 in Appendix displays a schematic

overview of stimuli. For even more detailed properties, see Vuust et al. (2006).

Experiment 2: The Microtiming Experiment

As for The Microtiming Experiment we generated 25 new groove excerpts lasting 30 seconds

each (see Figures A2-A6 in Appendix). Excerpts were recorded acoustically by professional

jazz musicians (JJH and the author JFS) on a standard drum kit and double (upright) bass

respectively, with MTL as a studio engineer. Drums and bass constitute the typical ‘rhythm

section’ in many contemporary musical genres, hence, most people should be used to pay

‘rhythmic attention’ to these instruments. It is also a constellation where microtiming

asynchronies may occur in groove contexts (Keil, 1987). Bass and kick drum are both located

low in the typical musical pitch range and time perception has indeed been found to be

generally superior when processing low compared to high musical pitch, as evidenced by

behavioural and brain activity measures (Hove, Marie, Bruce, & Trainor, 2014). Further, a

small pitch difference between instruments increases the chance that timing asynchrony is

discernible for the listener (Rasch, 1988). Finally, by only presenting bass and drum, both

playing a key role in timing perception, we simultaneously removed possible confounders.

The recorded groove excerpts were categorised into three levels (Low, Medium,

High) of what we called structural Complexity. Besides features of polyrhythm and

microtiming, syncopation is another common measure of structural complexity in a groove

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(Witek et al., 2014). Therefore, Complexity in this context refers to syncopation density as

well as the number of note onsets per bar (these two variables were confounded in the present

stimuli). The ‘low’ complexity excerpt (Figure A2 & A3) featured two notes per bar in the

bass, and no syncopations. ‘Medium’ (Figures A4 & A5) featured three bass notes per bar, in

which two of these were syncopated, and no drum syncopations. To be able to directly

examine the processing effect of a steady timekeeper, ‘low’ and ‘medium’ featured versions

with (Figures A3 & A5) and without (Figures A2 & A4) eighth notes hi-hat (‘HH’).

Generally, there is no obvious connection between an increase in event density caused by

steady timekeeping and structural complexity. Rather, as long the acoustical articulations of

the time-keeping instrument support rather than challenge the listener’s metric reference,

timekeeping will likely simplify the processing of the rhythmic material. ‘High HH’ (Figure

A6) was recorded in a single version only which included both hi-hat and snare drum. It

featured a more intricate drum kit pattern with two sixteenth note syncopations per bar in the

kick drum. The bass line was melodic and non-syncopated, ‘walking bass like’, with eight

eighth notes per bar. The structure was eight bars long, in contrast to the low and medium

complexity-grooves that had a repeated two-bar structure. Another difference between ‘high

HH’ and the other two complexity levels, was the fact that the former was inspired by a real

musical example, namely the R&B/soul-tune Really Love by the artist D’Angelo (2014),

where microtiming are in fact a part of the groove matrix between bass and drums. An

analysis performed by AD (with Amadeus Pro, version 2.3.1; see figure A7 in Appendix)

scrutinising the microtiming asynchrony between bass and drums in the original track of

‘Really Love’, showed that the timing of the electric bass is approximately timed 40-60 ms

behind the kick drum timing, at least in parts of the tune. This also enabled us to compare the

effects from an ‘ecological’ groove, with two ‘artificially made’ grooves (although all

excerpts were acoustically recorded by the author and his colleague on bass and drums,

respectively).

Table 1 Overview of stimuli in Experiment 2: The Microtiming Experiment

Note: *There was no ‘high’ condition, since the high complexity groove was presented with hi-hat only. Italicised cells contain names of conditions as used in text. All five groove-excerpts were presented in five microtiming conditions (relative placement of bass compared to drums): -80 ms, -40 m, 0 ms, +40 ms, +80ms.

Timekeeping:supportingmetricreference(eighthnotehi-hat)No Yes

Complexity: Low low(FigureA2) lowHH(FigureA3)Medium medium(FigureA4) mediumHH(FigureA5)High * highHH(FigureA6)

numberofsyncopations/noteeventsperbar

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All excerpts were 16 bars (in 4/4 bar) long, using a metronome click track set to 80

bpm as a common rhythmic grid. Eighty bpm is an adequate tempo for groove based music

(especially in the R&B /soul-genre; Danielsen et al., 2015), and also the actual tempo of

Really Love that served as the model for our high complexity groove. The notes played by the

bass player and drummer were not of identical duration and loudness throughout the

recordings, a result of intentionally trying to keep the stimuli as musical and natural as

possible. Bass and drum tracks were recorded separately and put together in Pro-Tools (with

help from technician MTL). Drums were recorded using one microphone on each drum (kick,

hi-hat, snare drum). Three microphones were also used for the double bass recording. Phase

alignment of the three bass tracks was performed, according to the timing from the contact

microphone – the ‘fastest’ of the three incoming signals. This signal happens artificially

early, compared to the normal acoustic properties of the double bass. To compensate for this,

the aligned bass tracks were moved 10 ms later/’to the right’. This was sufficient to give a

sense of bass and drums being together – ‘on the grid’. We used the ‘on the grid’ version as a

reference, and made four additional microtiming levels in terms of relative temporal timing

of the bass compared to the drum/metronome timing: bass 40 ms and 80 ms ahead (-) of, and

bass 40 ms and 80 ms after (+) the drums/metronome reference. In other words, there was a

stepof 40 ms between each level. All bass notes in each trial had the same relative temporal

position compared to the drum track, and the drum track itself had no internal microtiming

asynchronies. Each trial began with three metronome ticks, timed together with the drum set

recording. The last/fourth tick was removed to obscure which of the instruments that deviated

from the initial metronome reference.

In conclusion, Experiment 2 consisted of three grooves of different structural

complexity (varying degree of syncopations and note onsets per bar), in which the low- and

medium complexity groove were presented both with (‘HH’) and without a meter-supporting

time-keeping hi-hat. Each groove (‘low’, ‘low HH’, ‘medium’, ‘medium HH’, ‘high HH’)

were presented in five microtiming levels (including the on-the-grid version), making 25 the

total number of excerpts. See Table 1.

3.2.3 Setup and procedure Testing took place at the Cognitive Laboratory in the Department of Psychology, University

of Oslo, in a windowless and quiet room with constant illumination of about 170 Lux and

constant temperature and humidity. Remote Eye Tracking Device (RED) and I-View

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software commercially available from SensoMotoric Instruments (SMI), Berlin (Germany)

were used to record oculomotor data (pupil diameters, eye movements, fixations and blinks).

The RED system records quantitatively binocular pupil diameter with high precision

(detecting changes as a small as .0004 mm, according to SMI specs) using infrared light

lamps and an infrared light sensitive video camera. The sampling frequency was set low to 60

Hz. Instructions in English were given on a flat DELL LCD monitor with a screen resolution

of 1680 X 1050, installed above the RED video camera. The distance from the participant’s

eyes to the monitor and eye tracking device was set to 60 cm by use of a chin/head stabilizer.

Participants gave tapping responses during the active condition by pressing a key on the PC’s

keyboard (Dell L3OU) placed in front of the participant. The keyboard was linked to the PC

via a STLAB USB port hub. All participants wore a set of stereo headphones (Philips SBC

HP840) during the music listening.

All participants were individually tested by the author/experimenter who was present

in the room at all times with each participant. Most of the testing was done in the afternoon in

working days, occasionally in evenings. Both experiments were performed in each session.

Near-sighted participants were asked not to use their glasses, and female participants to

refrain from using mascara, since these are often source of artefacts during eye tracking. All

participants were requested to keep their eyes open (in order to obtain continuous pupil

recordings), except for short eye blinks, and to maintain gaze on a black fixation cross (+)

displayed in the middle of the monitor. Furthermore, they were informed that it was possible

to rest their eyes when the fixation cross was not present. The fixation cross was used to

facilitate anchoring gaze and avoid that participants would look away from the screen;

however, it was not necessary for collecting pupil sizes. A neutral grey colour (pixels’ RGB:

163,163,163 for the Experiments, and RGB: 166,166,166 for MET) was used as background.

All experiment sessions began with a standard 5-point eye calibration procedure, a general

instruction slide, and individual adjusting of the sound volume. Each trial was self-paced, i.e.,

initiated by the participant by pressing the spacebar. Three seconds before and during the

music clips, the fixation cross was displayed in the middle of the screen. Each participant was

first exposed to the 25 microtiming excerpts in a ‘listening only’ condition. The order of

stimuli was pseudorandomized, with the constraint that it was made sure that none same

groove types (e.g., ‘low HH’) were played back-to-back. The pseudorandomized order of

stimuli was counterbalanced so that half of the participants in each group (musicians/non-

musicians) began with one of the two blocks. After completing the listening of each excerpt,

the participants were asked two questions (on a 5-step Likert scale: 1 = not at all, 5 = to a

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great extent): 1) To what extent does listening to this rhythm, make you want to move your

body? and 2) To what extent do you find that this rhythm sounds musically well-formed?

Question 1 tapped onto the concept of groove experience, treating groove exclusively as a

sensorimotor phenomenon, according to Madison’s (2006) definition of groove. We did not

enquire about ‘positive affect’, since microtiming stimuli were brief bass and drums-tracks,

therefore not ‘ecological’ musical clips; hence we did expect them to have relatively low

aesthetic value per se. Question 2 enquired about rating of musical ‘well-formedness’, similar

to Davies’ et al., (2013) ‘musical naturalness’. After listening to all microtiming stimuli, the

participants were exposed to Experiment 1, i.e., the polyrhythmic stimuli and the

corresponding control stimuli (counterbalanced for half of the participants). After a break,

there was an introductory test-trial where participants were asked to tap with their index

finger, synchronising to a 30 second isochronous metronome beat. After this, participants got

the following instruction:

The clips you will hear from now on will begin with the metronome pulse, then the

music will start. Tap with the metronome and, as steady and regularly as you can,

continue tapping the basic pulse of the music, until the music stops.

If participants didn’t perform this task as instructed, the experimenter alerted verbally the

participant, repeating the instructions (e.g., ‘only tap the basic pulse given by the metronome,

not any rhythmic pattern’). Following this, the polyrhythmic and microtiming stimuli were

presented again in an active tapping condition; however, in reverse order for all participants.

After completing the tapping condition, participants were given the Musical Ear Test

(‘rhythm part’ only). The test began with instructions and two practice trials. Every new trial

was initiated by pressing the spacebar. All participants got the MET stimuli in the same order

and no feedback was given during testing. Eye-tracking data was also recorded while

participants did MET, but these data were not analysed. Table A1 in the Appendix shows an

overview of the complete experimental procedure. Immediately following the experiment

session (taking approximately 70 min to complete), the participant filled out an online

questionnaire. The questionnaire was based on the original form from the study of Laeng et

al. (2016), but adapted to the present study.

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3.2.4 Data processing Pupil, tapping and subjective rating data were exported from BeGaze software (SMI, v.3.4)

and imported to MicrosoftÒ Excel where they were sorted and categorized. As pupil sizes of

left and right eye were identical, participants having maintained fixation on a central cross,

no mean value of the two eyes was necessary. Subtractive baseline correction in different

variations are commonly applied in pupillometry research (e.g., Bianchi, Santurette, Wendt,

& Dau, 2016; Laeng & Sulutvedt, 2014). In the present study, average individual left eye’s

pupil size by each trial (lasting 30,000 ms) were baseline-corrected by the average pupil size

taken 3000 ms before stimulus onset for each participant per trial, giving a measure of ‘pupil

size change’ (corrected for luminance and expressing dilations from baseline as positive

numbers and relative constrictions as negative numbers).

As for the tapping data, the timing of each tap was compared to a corresponding

reference point of the musical pulse, the latter operationalized as X milliseconds after

stimulus onset. In Experiment 1, the inter-pulse-interval for musical reference points was 500

ms (120 bpm), while for Experiment 2 it was 750 ms (80 bpm). The timing of the

drums/metronome (not the double bass) served as the basis for the reference points in the

microtiming stimuli. The first eight taps in each trial in both experiments were excluded from

the analyses. To identify and exclude outliers, pre-processing of tapping data was done by

custom scripting in MATLAB (performed by KN), according to the following rules:

• Exclude (and count) double taps, defined as more than one tap within a single time

period, corresponding to the time period of ±50% between one reference point and the

next/previous reference point (±250 ms around each reference point when inter-pulse-

interval was 500 ms, and ±375ms when inter-pulse-interval was 750 ms).

• Count missing reference points, defined as reference points without any taps within the

time period described above.

• Exclude (and count) complete trials that have more than six reference points containing

double or no taps.

The typical dependent variables in tapping studies are either ‘absolute’ offsets

between reference points and the tap and their standard deviation; additionally, one can

compute the mean and standard deviation of intertap intervals (ITI; Repp & Su, 2013). From

preliminary tapping analyses, we discovered the presence of an artifactual ‘time lag’ or

‘instability’ in the software system (possibly generated from the use of a standard keyboard

and USB port in collecting taps; as confirmed by SMI’s personal communication), that made

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the absolute offsets non-reliable and possibly non-coherent between participants. For

example, when tapping to the ‘plain condition’ of The Polyrhythm Experiment, professional

musicians average mean offset from reference points were +109 ms, with an inter-participant

variance (SD) of 33 ms. Both measures far exceeded what we could expect from tapping

behaviour in the musician group according to previous studies (e.g., Repp & Su, 2013). This

artefact prevented us from analysing the absolute offsets, which could have been useful in

exploring how microtiming influenced absolute placements of taps. Nevertheless, the intra-

participant tapping offset distributions appeared reliable and within the normal range (the

mean SD offset when musicians tapped to ‘the plain condition’ of The Polyrhythm

Experiment was 18 ms). We therefore selected ‘SD offset’ data as the main dependent

variable for tapping in our analyses, used as a reversed measure of ‘tapping accuracy’. There

was still, however, a small element of uncertainty relating to exclusions of occasional taps: as

discussed above, taps were included when being ±50% around each reference points (±250

ms in Experiment 1 and ±375 ms in Experiment 2). Since taps generally seemed to be placed

±100 ms too late due to technical issues, this meant that ±150ms/±275ms-taps in Experiment

1 and 2, respectively, were considered ‘outliers’ and excluded from the statistical analyses.

We decided to ignore the intertap interval (ITI), since missing and excluded double taps

tended to generate double ITIs (for example 1000 ms instead of 500 ms), as corresponding

reference points did not have any attached taps.

Missing pupil data were generally very few (1.3% or 23 out of totally 1728 trials

across both experiments); thus, there was no need to exclude any participant from the pupil

analyses due to data loss. Tables A2-A5 in Appendix summarise tapping data exclusions,

sorted into three categories for musicians and non-musicians, respectively: 1) number of

excluded trials (having more than six reference points containing double or missing taps); 2)

number of reference points without taps in included trials; 3) number of double taps in

included trials. Across the two experiments, 8.7% or 75 out of totally 864 tapping trials were

excluded, however, all participants’ valid trials were included in our analyses. Group

differences in tapping exclusions were visually scrutinised, but not statistically analysed. As

for the repeated-measures ANOVAs, missing trials in both pupil and tapping data were

estimated using that participant’s group (musician/non-musician) mean pupil change/SD

offset in the respective condition so as to fill out missing cells in the statistical spreadsheet.

Statistical analyses were performed with Statview and/or SPSS (v.24) software.

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4 Results 4.1 Music Engagement Questionnaire

From the questionnaire (with 5-step Likert scales), we found from independent samples t-

tests that musicians enjoyed more listening to music in general (M = 5; SD = 0) than non-

musicians (M = 4.63; SD = .62); t(30) = 2.42, p < .001. Musicians and non-musicians did not,

however, differ significantly in how interesting they found participating in the present study,

or how interesting they found the study’s musical excerpts.

4.2 A general pupillary drifting effect Figure 4 shows participants’ mean pupil diameter size (in mm) as a function of time (0-30

seconds) for all trials in both experiments combined. By visual inspection we can observe a

response peak 5-7 seconds after stimulus onset, and later a negative ‘drifting’ effect with a

more or less continuous decay – the pupil generally tended to decrease in size as the trials

progressed. Hence, it was reasonable to expect the mean baseline-corrected pupil sizes in

both experiments to be either close to or below the zero level, as the baseline-condition was

only 3 seconds in duration and measured right before each stimulus, whereas the

experimental condition took 30 seconds.

Figure 4. Mean pupil size as function of time (s) for all experimental trials combined. Error bars: ±1 Standard error(s).

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4.3 Experiment 1: The Polyrhythm Experiment 4.3.1 Pupil responses

A 2*2*2 mixed between-within repeated-measures ANOVA of mean baseline-corrected

pupil change dependent on Musicianship (musicians vs. non-musicians), Rhythm (‘the

polyrhythmic condition’ vs. ‘the plain condition’) and Activity (‘passive’ [listen only] vs.

‘active’ [tapping with the beat]), was performed. ‘The polyrhythmic condition’ yielded larger

pupil dilation change (M = .06 mm; SD = .29) than ‘the plain condition’ (M = -.03 mm; SD =

.26), approaching a significant difference, F(1, 30) = 4.0, p = .054, in the predicted direction.

In addition, we found a significant, F(1, 30) = 35.89, p < .001, difference between the active

(M = .12 mm; SD = .27) and passive (M = -.09 mm; SD = .24) conditions, indicating that

tapping the beat while listening, triggered a larger pupil dilation change than listening only.

However, there was no significant pupil change difference between musicians and non-

musicians. No two- or three-way interactions were found.

4.3.2 Tapping accuracy Musicians had fewer excluded participants,

double taps and missing reference points in

both the ‘polyrhythmic’ and ‘plain’

conditions (Table A2 in Appendix). When

only included taps and trials were taken into

account, a mixed between-within repeated-

measures ANOVA of tapping accuracy (SD

offset) was performed. Independent variables

were the same as in the pupil response

analysis (except Activity, since the active

condition was the only tapping condition). As

expected, tapping accuracy was significantly

lower (i.e., higher SD offset) when tapping

the main meter in ‘the polyrhythmic

condition’ (M = 64.09 ms; SD = 40.22),

compared to tapping the main meter in ‘the

plain condition’ (M = 27.28 ms; SD = 16.12), F(1, 30) = 37.45, p < .001. Also, a significant

effect of Musicianship, F(1, 30) = 28.86, p < .001, appeared. Specifically, the musicians’

Figure 5. Tapping accuracy (SD offset in ms; low values represent higher accuracy) for musicians and non-musicians when tapping the main meter in the ‘plain’ and ‘polyrhythmic’ conditions. Error bars: ±1 Standard error(s).

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tapping accuracy (M = 28.68 ms; SD = 24.77) exceeded the non-musicians’ (M = 62.69 ms;

SD = 36.96). In addition to the main effects, the expected interaction effect of Rhythm with

Musicianship was significant, F(1, 30) = 5.32, p= .028, as shown in Figure 5, where the

relative difference in tapping performance between musicians and non-musicians were

stronger with cross-rhythms, compared to the plain condition.

4.4 Experiment 2: The Microtiming Experiment 4.4.1 Pupil responses

A 2*5*5*2 mixed between-within

repeated-measures ANOVA of mean

baseline-corrected pupil change

dependent on Musicianship (musicians

vs. non-musicians), Microtiming

asynchrony (-80 ms, -40 ms, 0, +40 ms,

+80 ms), Complexity (‘low’, ‘low HH’,

‘medium’, ‘medium HH’, ‘high HH’)

and Activity (‘passive’ vs. ‘active

conditions’, was performed.

The effect of Musicianship.

As in Experiment 1, there was no

significant pupillary response difference

between musicians and non-musicians,

which was also the case when we

performed a separate analysis where the microtiming asynchronies from zero were collapsed

over direction into three levels (0 ms, ±40 ms and ±80 ms). The analysis did not show any

interaction effects of musicianship by any experimental condition.

The effect of Activity.

Pupil dilation responses were, as in Experiment 1, significantly stronger when

participants tapped to the beat (M = -.02 mm; SD = .26), compared to when they listened only

(M = -.19 mm; SD = .26), F(1, 30) = 82.3, p < .001.

The effects of Microtiming asynchrony.

A significant main effect of Microtiming asynchrony magnitudes on pupils change

emerged, F(4, 120) = 6.08, p < .001, as shown in Figure 6. There was an increased pupil

Figure 6. Pupil change as a function of Microtiming asynchrony magnitudes between bass and drums for both groups. Error bars: ±1 standard errors.

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response as microtiming asynchrony increased in either direction (i.e. bass timed before

drums [-] or drums timed before bass [+]). Pairwise contrasts (see Table 2) revealed that the 0

ms condition yielded significantly less pupil dilation change than the -80 ms, +80 ms and +40

ms microtiming asynchrony conditions. The 0 ms and -40 ms conditions, did not generate

significantly different pupil responses. The response difference between the -80 ms and -40

ms conditions was statistically significant, while this was not the case between the +40 ms

and +80 ms conditions. The latter indicated that in the present experiment, changing the

microtiming asynchrony from 40 ms to 80 ms demanded more effort in the perceptual system

when bass was timed before drums, than when drums were timed before bass. No other

significant differences between the five Microtiming conditions were found.

Table 2 Pairwise contrasts for main effects of Microtiming and Complexity on pupil change

Note: a negative contrast value indicates that the mean for the condition in the leftmost column is lower than for the condition to the right. Statistically significant (p < .01) contrasts in boldface.

The mixed ANOVA also revealed a statistically significant Activity by Microtiming

asynchrony interaction, F(4, 120) = 9.78, p < .001. This could be explained by no effect of

Activity for the extreme -80 ms condition, which seemed to create approximately the same

mental effort independent of activity.

The effects of Complexity.

No main effect of Complexity (i.e., different levels of syncopations/note onsets per

bar) on pupil change was found. Unsurprisingly, a separate 2*3*5*2 repeated-measures

ANOVA that compared pupil change from the three complexity levels only, that included hi-

hat (‘Low HH’, ‘Medium HH’, ‘High HH’), also failed to reveal an effect. We did, however,

find a three-way interaction between Microtiming, Complexity, and Passive/Active, F(16,

480) = 1.71, p = .042, that was difficult to interpret and most likely reflecting spurious effects

resulting from the small number of trials constituting each condition. All other main and

interaction effects were weak and statistically non-significant, including the effect of

PairwisecontrastsformaineffectsofMicrotimingandComplexityMicrotiming:

Contrast SE p Contrast SE p-80 -40 0.045 0.016 0.010 low lowHH -0.005 0.018 0.768-80 0 0.070 0.019 0.001 low medium -0.031 0.016 0.062-80 +40 0.025 0.016 0.127 low mediumHH -0.005 0.017 0.754-80 +80 0.014 0.015 0.371 low highHH -0.020 0.020 0.336-40 0 0.026 0.016 0.132 lowHH medium -0.025 0.015 0.100-40 +40 -0.019 0.013 0.131 lowHH mediumHH 0.000 0.018 0.999-40 +80 -0.031 0.015 0.057 lowHH highHH -0.014 0.015 0.3400 +40 -0.045 0.015 0.006 medium mediumHH 0.025 0.011 0.0330 +80 -0.056 0.016 0.002 medium highHH 0.011 0.015 0.470+40 +80 -0.011 0.013 0.407 mediumHH highHH -0.014 0.016 0.382

Complexity:

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‘Timekeeping’ (comparing the ‘low’ and ‘medium’ complexity grooves with and without hi-

hat, respectively) on pupil response, investigated in a separate ANOVA.

4.4.2 Tapping accuracy Across all conditions of Experiment 2, musicians had fewer double taps but more missing

reference points in included trials than the non-musicians (Tables A3-A5 in Appendix). Yet,

it is probable that non-musicians had more missing reference points, on the basis of having

far more excluded trials (63) than the musicians (5). Note that excluded trials cover a

substantial amount of missing reference points. Taking into account only included trials and

taps, we performed a 2*5*5 mixed between-within ANOVA of SD offset, with the same

independent variables as in the pupil data analysis above (except Activity, since the active

condition was the only tapping condition).

The effect of Musicianship.

The analysis revealed a main effect of Musicianship, F(4, 120) = 45.97, p < .001.

Musicians (M = 23.11 ms; SD = 11.50) outperformed non-musicians (M = 80.08 ms; SD =

54.64) in tapping accuracy across all levels of microtiming asynchrony and complexity, as

can be seen in both graphs of Figure 7.

Figure 7. Tapping accuracy (SD offset in ms), for musicians and non-musicians as a function of Microtiming asynchrony (right) and structural Complexity (left). Error bars: ±1 standard errors. The effects of Microtiming asynchrony.

The main effect of Microtiming asynchrony was statistically significant, F(4, 120) =

10.49, p < .001. However, the analysis also revealed a statistically significant Microtiming

asynchrony by Musicianship interaction, F(4, 120) = 5.77, p < .001. The left side graph of

Figure 7 shows that birectional microtiming asynchrony decreased tapping accuracy;

especially the -80 ms condition seemed to have a negative effect. This was confirmed by

0

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Table 3 showing the results for pairwise contrasts for the main effect of Microtiming

asynchrony, for non-musicians and musicians separately. For the musicians, in fact, it was

only the -80 ms condition that significantly increased SD offset compared to the other

conditions, but for the non-musicians, also the +80 ms conditon had a significant negative

sensorimotor synchronisation effect (compared to the -40 ms condition). The -40 ms and +40

ms conditions did generally not decrease tapping accuracy compared to the on-the-grid-

excerpts.

Table 3 Pairwise contrast for the effect of Microtiming asynchrony on tapping accuracy (SD offset) – overall and separately for non-musicians and musicians.

Note: a negative contrast value indicates that the mean for the condition in the leftmost column is lower than for the condition to the right. Statistically significant (p < .01) contrasts in boldface.

The effects of Complexity.

The mixed ANOVA revealed a main effect of Complexity, F(4, 120) = 5.76, p < .001.

From the right side graph of Figure 7 it is evident that this main effect was primarily due to a

positive influence of Timekeeping (hi-hat eighth notes vs. no hi-hat eighth notes). That is,

participants’ tapping accuracy was significantly higher (lower SD Offset) in both the low and

medium-grooves with hi-hat, compared to the low and medium-grooves without hi-hat. The

latter is displayed in Table 4, showing the pairwise contrasts for the main effect of

Complexity, and for non-musicians and musicians separately.

To control for (exclude) the effect of Timekeeping, we comparing tapping accuracy in

the two groups on the grooves with hi-hat only (‘low HH’, ‘medium HH’ and ‘high HH’). A

2*3 mixed between-within ANOVA with Musicianship and this simplified Complexity

variable as independent variables, gave a significant interaction effect of Complexity with

Musicianship, F(2, 60) = 3.31, p = .043, in addition to the expected main effects. Although

Figure 7 displays all five complexity conditions, we can see from the graph that both groups’

tapping accuracy decreased as Complexity (number of syncopations/sound events per bar)

increased, but to a greater degree in the non-musicians; hence the interaction effect is mainly

Microtiming: Non-musicians. Musicians.Contrast SE p Contrast SE p Contrast SE p

-80 -40 19.537 3.763 0.000 34.266 7.450 0.000 4.809 1.074 0.000-80 0 15.472 4.503 0.002 24.532 8.869 0.014 6.413 1.569 0.001-80 +40 15.696 4.366 0.001 25.521 8.670 0.010 5.871 1.044 0.000-80 +80 7.831 3.309 0.025 11.137 6.535 0.109 4.524 1.050 0.001-40 0 -4.065 2.601 0.129 -9.734 5.047 0.073 1.604 1.264 0.224-40 +40 -3.841 3.199 0.239 -8.745 6.314 0.186 1.063 1.031 0.319-40 +80 -11.706 2.797 0.000 -23.128 5.442 0.001 -0.285 1.295 0.8290 +40 0.224 2.895 0.939 0.989 5.683 0.864 -0.541 1.106 0.6320 +80 -7.642 2.922 0.014 -13.395 5.764 0.035 -1.889 0.963 0.069+40 +80 -7.865 3.140 0.018 -14.383 6.220 0.035 -1.347 0.871 0.143

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driven by them. However, the variance of the non-musicians’ tapping data was much larger

than the variance of the musicians’ tapping data, which might be a reason for why no

significant tapping accuracy differences occurred between any of the three complexity levels

(‘low HH’, ‘medium HH’ and ‘high HH’) when alpha was set low to .01 to control for

multiple comparisons.

Table 4 Pairwise contrast for the effect of Complexity on tapping accuracy (SD offset) – overall and separately for non-musicians and musicians.

Note: a negative contrast value indicates that the mean for the condition in the leftmost column is lower than for the condition to the right. Statistically significant (p < .01) contrasts in boldface.

To summarise the main findings from tapping data in Experiment 2: musicians kept

time more accurately than non-musicians; microtiming asynchrony (though only the two

extreme conditions, ±80 ms) negatively influenced tapping accuracy in both groups, but non-

musicians to a substantially greater degree than musicians. Tapping accuracy generally

tended to be higher in the low complexity than the high complexity grooves. Finally,

timekeeping (i.e., presence of meter-supporting hi-hat eighth notes) increased tapping

accuracy in both groups.

4.4.3 Rating scales The two rating scales were based on the following questions: 1) To what extent does listening

to this rhythm, make you want to move your body? (‘move body’) and 2) To what extent do

you find that this rhythm sounds musically well-formed? (‘well-formed’) Participants’

responses on the two scales were moderately to highly correlated (non-musicians: r = .56, p =

.000; musicians: r = .80, p < .001). We performed a 2*5*5*2 mixed between-within ANOVA

of subjective ratings, dependent on Musicianship (musicians vs. non-musicians), Microtiming

asynchrony (-80 ms, -40 ms, 0, +40 ms, +80 ms), Complexity (‘low’, ‘low HH’, ‘medium’,

‘medium HH’, ‘high HH’) and Rating Scales (‘move body’ vs ‘well-formed’).

Complexity: Non-musicians. Musicians.Contrast SE p Contrast SE p Contrast SE p

low lowHH 13.704 3.474 0.000 15.307 6.729 0.038 12.101 1.730 0.000low medium -0.087 3.679 0.981 -1.054 7.092 0.884 0.880 1.962 0.660low mediumHH 13.232 3.437 0.001 17.063 6.728 0.023 9.402 1.414 0.000low highHH 1.645 5.072 0.748 -5.888 10.085 0.568 9.178 1.103 0.000lowHH medium -13.791 3.295 0.000 -16.361 6.339 0.021 -11.221 1.800 0.000lowHH mediumHH -0.471 2.539 0.854 1.756 4.894 0.725 -2.699 1.352 0.064lowHH highHH -12.059 5.304 0.030 -21.195 10.560 0.063 -2.923 1.020 0.012medium mediumHH 13.319 2.768 0.000 18.117 5.410 0.004 8.522 1.174 0.000medium highHH 1.732 5.396 0.750 -4.834 10.638 0.656 8.298 1.815 0.000mediumHH highHH -11.587 5.580 0.047 -22.951 11.098 0.056 -0.224 1.178 0.852

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The effect of Musicianship.

We found a significant interaction effect of Microtiming by Musicianship, F(4, 120) =

10.25, p < .001, indicating that musicians were more responsive to microtiming than non-

musicians, using a wider range of the rating scale. As shown in Figure 9, musicians rated the

conditions with largest microtiming (±80 ms) lower, and the on-the-grid condition higher

than the non-musicians.

The effects of Microtiming asynchrony.

The main effect of Microtiming, F(4, 120) = 30.06, p < .001, provides evidence that

bidirectional magnitudes of microtiming asynchrony were negatively related to subjective

rating on both rating scales. In other words, when ratings were higher, the closer aligned the

bass and drums were timed (see Table 5). Furthermore, conditions where bass was timed

after the drums, were preferred in favour of (i.e., rated higher than) the conditions where bass

was timed ahead of the drums. As already mentioned, Figure 9 clearly shows that the

musicians are to a greater degree responsible for this trend. Also, a significant interaction

effect of Rating scales by Microtiming, F(4, 120) = 5.61, p < .001, emerged. From Figure 10,

we observe that participants rated excerpts higher on ‘well-formed’ than ‘move body’, and

that this difference tended to be larger with less bidirectional microtiming asynchrony.

Figure 9. Interaction effect of Musicianship by Microtiming asynchrony on subjective ratings (mean of both scales). Error Bars: ±1 Standard Error(s).

Figure 10. Interaction effect of Rating scales by Microtiming asynchrony, on subjective ratings. Error Bars: ±1 Standard Error(s).

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Table 5 Pairwise contrasts for main effects of Microtiming asynchrony and Complexity on subjective ratings.

Note: a negative contrast value indicates that the mean for the condition in the leftmost column is lower than for the condition to the right. Statistically significant (p < .01) contrasts in boldface.

The effects of Complexity

As expected, subjective ratings increased as Complexity also increased, F(1, 30) =

27.04, p < .001. Pairwise contrasts for all conditions are presented in Table 5.

In addition, we found a positive ‘Timekeeper effect’ on subjective ratings. Low and

medium complexity grooves with hi-hat were rated higher on both scales than the

corresponding grooves without hi-hat. On ‘Wanting to move’ (with HH: M = 2.53, without

HH: M = 2.1), t(31) = 5.32, p < .001. On ‘Musically well-formed’: (with HH: M = 3.04,

without HH: M = 2.71), t(31) = 4.70, p < .001. This effect is also evident from the pairwise

contrasts in Table 5, where the two

rating scales are combined.

Finally, from the initial

ANOVA-analysis we found a

Microtiming asynchrony by

Complexity interaction effect on

subjective ratings, F(16, 480) =

3.78, p < .001. We can appreciate

from Figure 11 that microtiming

asynchronies seemed to give larger

fluctuations in ratings (steeper

profiles on the graph) as groove

complexity increased.

Microtiming: Complexity:Contrast SE p Contrast SE p

-80 -40 -0.616 0.105 0.000 Low LowHH -0.272 0.075 0.001-80 0 -0.931 0.130 0.000 Low Medium -0.566 0.070 0.000-80 40 -0.828 0.108 0.000 Low MediumHH -0.616 0.095 0.000-80 80 -0.341 0.101 0.002 Low HighHH -1.122 0.151 0.000-40 0 -0.316 0.077 0.000 LowHH Medium -0.294 0.080 0.001-40 40 -0.212 0.080 0.013 LowHH MediumHH -0.344 0.088 0.001-40 80 0.275 0.094 0.006 LowHH HighHH -0.850 0.153 0.0000 40 0.103 0.059 0.093 Medium MediumHH -0.050 0.076 0.5160 80 0.591 0.115 0.000 Medium HighHH -0.556 0.145 0.00140 80 0.487 0.088 0.000 MediumHH HighHH -0.506 0.154 0.003

Figure 11. Interaction effect of groove Complexity and Microtiming asynchrony on subjective ratings (based on two rating scales, ranging 1-5). Error Bars: ±1 Standard Error(s).

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5 General Discussion 5.1 General effects on pupil responses and tapping

accuracy support DAT and PC The results from both experiments on pupil and tapping response strongly support the notion

that the brain allocates increased effort to process musical stimuli whenever rhythmic input

challenges fundamental metrical models. We have little foundation, however, to conclude

whether either PC or DAT better explain the data, also because it remains unclear as to

whether the two accounts can really generate diverging predictions (at least in the present

context). Looking at pupil responses first, Experiment 1 shows that both listening to, and

tapping the main meter (M) in a rhythm containing a competing 4:3 pulse train (C), yielded

higher measures of effort for both groups, as indexed by pupillary change, compared to

listening to and tapping the main meter in a comparable non-polyrhythmic context. The

above findings fit the hypotheses given by DAT and PC, since according to these accounts,

an acoustically-salient counter-rhythmic (C) pulse level will challenge the participants’

maintenance of the neural oscillations (DAT)/predictive models (PC), which represent the

original but acoustically-absent main meter level (M). Like our results suggest, this demand

involves mental processing power. Similarly, in Experiment 2, when microtiming

asynchronies between bass and drums in the groove excerpts emerged and increased, a

corresponding increase in cognitive workload reflected in the pupil was accordingly

observed. Based on the theories, increased attentional processing demands would be a result

of participants experiencing prediction error, and adjusting the neural oscillations phase

and/or widening the temporal attentional focus (DAT)/updating the anticipated mental model

(PC), so as to account for the incoming rhythmic events across the 30-second music duration.

The larger the asynchronies between the timing of the bass and drums, the more the need for

interval adjustments; interestingly, we did measure a higher degree of effort concomitantly to

level of asynchrony. One should note that the pupil changes may also be indicative of

response conflicts (Kamp & Donchin, 2015). The inherent temporal ambiguity of exact beat

placement that emerged, especially in the most ‘extreme’ microtiming asynchrony

conditions, may have generated response conflicts as to mentally or physically lay down the

tap. Indeed, the ±80 ms conditions may also have caused metric uncertainty, since it

approached the perceptual threshold for temporal integration; that is, when microtiming

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asynchrony exceeds 100 ms, sound onsets – even when in a similar frequency range – tend to

be experienced as separate, rather than integrated events (Repp, 2005; Warren, 1993).

General tapping results did partly mirror pupil data and appeared to be consistent with

expectations. We found that both polyrhythm and bidirectional (bass ahead of drums vs.

drums ahead of bass) microtiming asynchrony influenced tapping accuracy negatively for

both groups. However, only when exposed to the ±80 ms microtiming asynchrony conditions,

did the accuracy significantly decrease from the on-the-grid level; the ±40 ms conditions did

not disturb participants’ fine grained experience of beat placement to the degree that it was

reflected in tapping accuracy. This divergence might be explained by the 80 ms, but not 40

ms asynchrony, approaching the threshold for temporal integration, as mentioned above.

Furthermore, we generally found a higher effort in the active ‘tap with the beat’

condition, compared to the passive ‘listen only’ condition. This is not surprising, as music

production is clearly a more complex activity than music perception (Zatorre et al., 2007).

Tapping involves an active participation in the groove that demands a continuous and

coordinated motoric ‘realization’ of the internal metric model. Additionally, making an

explicit response may enhance the intensity of attentional focusing (Moresi et al., 2008; Van

der Molen, Boomsma, Jennings, & Nieuwboer, 1989) over passively listening, which would

also be expected to yield larger pupil sizes in the former than the latter. Finally, we found an

overall negative ‘drifting’ of the pupil sizes from around 5-7 seconds, and throughout the 30

seconds of the trial length. This is not unusual with pupillometric measurements and may be

interpreted as a decrease in general arousal levels with protracted focus on a task. However,

given the specific predictions generated by our two cognitive meter perception theories, the

response peak followed by gradual and continuous decay may also be interpreted as a

correspondingly gradually decreasing anticipation error, resulting from the participants’

mental models becoming initially challenged, but then increasingly adjusted, as participants

were accustomed to the rhythms.

5.2 Effects of musical expertise Numerable studies have documented the enhanced perceptual skills of musicians for rhythm

and meter (e.g., Chen et al., 2008; Drake et al., 2000; Geiser et al., 2010; Matthews et al.,

2016; Stupacher et al., 2017). The rhythm part of Musical Ear Test did also distinguish the

groups, as musicians outperformed the non-musicians. Both groups however, performed

slightly better than earlier samples of musicians and non-musicians (Wallentin et al., 2010).

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This could be due to indeed more rhythmic competent participants selected for the present

study or due to the test situation; that is, our participants were permitted short breaks between

each trial and initiated trials themselves by pressing the spacebar. This was in contrast to the

Wallentin et al. (2010) study, where the test ran as a whole without any stops, with only one

‘resting’ bar between each trial.

It was of great interest to us to investigate the effect of musical expertise on the

cognitive processing of polyrhythm and microtiming especially. In the following, we will

look at several explanations that can account for the lack of pupillary group differences found

across both experiments, as well as discussing the clear-cut and expected effects of

musicianship on tapping data.

5.2.1 Pupillary response was unaffected by musical expertise As previously mentioned, MMN-studies have often found stronger brain responses in

musicians than in non-musicians, for example to instances of harmonic (Brattico et al., 2008)

and metrical incongruence (Vuust et al., 2005) and even syncopations (which musicians

clearly should be more accustomed to; Vuust et al., 2009). Indeed, enhanced perceptual

sensitivity is supported in part by increased involvement of the working memory system

(musical working memory is also the target variable of Musical Ear Test). Chen et al. (2008)

found an increased activation of the dorsolateral prefrontal cortex (being associated with

working memory function) in musicians compared to non-musicians when processing

meter/rhythmic structures. Therefore, working memory recruitments imply heightened

processing demands which should also be reflected in measures of pupillary changes (Laeng

et al., 2011). Based on such reasoning and the MMN-results, it was expected that musicians

displayed increased pupillary response to rhythmic complexity. However, in other lines of

research, expertise has been negatively related to mental effort both in musical tasks (Bianchi

et al., 2016) and other domains, like face recognition memory (e.g., Wu, Laeng, &

Magnussen, 2012), indicating a more efficient processing of relevant stimuli in experts than

in novices. Thus, although we generally expected some pupillary response effects of

musicianship, the direction (decreased or increased response) was less certain. Given that, in

fact, our analyses failed to reveal any significant differences, it remains unclear what effects

expertise has on effort in our specific domain.

Two recent studies, Bouwer et al., (2014) and Damsma & van Rijn (2017), may partly

inform this discussion. Both investigations came to the conclusion that basic meter perception

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is independent of musical expertise, since they found no effect of musical expertise on MMN

and on pupil response respectively, when omitting beats placed at metrically different spots

in ‘simple’/basic rhythm excerpts. This may have implications for the lack of group pupil

differences in our study during processing of microtiming. Given the notion that the internal

metric model strength for musicians and non-musicians is indeed similar for ‘basic’ rhythms

implying a ‘simple meter’, and the grooves of the present study are in the domain of ‘basic’

(despite including micro-metrical ambiguities), it follows that general enculturation, shared

by both groups, would render possible an ‘adequate’ processing of these stimuli. However,

this account apparently contrasts with the clear-cut difference observed between our groups

in tapping performance. Moreover, remaining unclear is the absence of pupillary effects

between groups when participants processed cross-rhythms, since such rhythms are more

‘complex’ than ‘basic’. Indeed, group differences related to polyrhythmic processing should

be expected based on earlier studies’ inverse correlations between rhythmic expertise in

professional musicians and brain activity in the area of BA47 (related to the processing of

metric structures) when listening to 4:3 polyrhythm (Vuust et al., 2006; Vuust, Wallentin,

Mouridsen, Østergaard, & Roepstorff, 2011). However, a very recent study by Stupacher et

al., (2017) suggests an alternative interpretation since they failed to find any effect of musical

expertise on EEG response when participants listened to a 3:4 cross-rhythmic sequence.

However, the predicted group difference in persistent endogenous oscillations did emerge

during the silent periods of 2-3 seconds that immediately followed each polyrhythmic

sequence. The authors speculated that under music listening, measurable brain activity

constitute a resultant of the close interaction between bottom-up (rhythm) and top-down

(meter) perceptual processes. Bottom-up responses are more or less hardwired, probably

independent of musical expertise, and activated independently of which metric framework

that is endorsed while listening. Top-down mechanisms in contrast, represents the metric

mental model. Importantly, neural oscillations generated from bottom-up processing were

found to have stronger amplitudes than top-down processes (Stupacher et al., 2017). Hence,

under music listening, expertise-sensitive top-down mechanisms could have been masked by

the strong effect of bottom-up rhythm perception, at least when measuring domain general

brain activity (pupil dilation, EEG), not activity in specific areas (i.e., BA47). In the silent

period, however, bottom-up processing is removed. Unfortunately, we did not record pupil

sizes within the silent periods following the auditory stimuli.

An additional account is the possibility that musicians have a higher initial response (i.e.,

prediction error) to metric challengers, as MMN-studies typically have suggested, but they

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can also adjust their internal models more efficiently (using less effort) over a period of time,

so that the averaged pupil dilation over the 30 seconds would appear similar to the non-

musicians’. Furthermore, pupil response can index aesthetic experiences in music listening

(Laeng et al., 2016), which may provide yet an alternative account for the lack of pupillary

group differences in The Polyrhythm Experiment especially. Because musicians and non-

musicians did rate the experiments’ musical stimuli to be equally interesting, it is not

unreasonable to also expect similar ‘aesthetic’ pupillary response from the two groups, when

being exposed to more and perhaps also more interesting musical information (i.e., in ‘the

polyrhythmic condition’).

One obvious explanation for the negative finding across both experiments is statistical

power. In other words, there might be a difference between groups in the present sample, but

that N is too low to be able to confirm it. After all, 16 participants constitute a small group in

general for psychology experiments as well as pupillometry experiments. More participants

might have brought to surface ‘hidden’ group differences. We might also argue that possible

expertise effects on effort were obscured because both our groups were to some extent

musically skilled, as the non-musicians’ scores on Musical Ear Test (MET) suggested, being

higher than the score of earlier samples of non-musicians (Hansen et al., 2012; Wallentin et

al., 2010). Two non-musician participants even scored higher on MET than the professional

jazz musicians with the lowest scores. In fact, the non-musician group included both non-

musicians and amateur musicians. Had the non-musicians group been singled out in two

separate groups (non-musicians vs. ‘amateur’ musicians), this may also have influenced the

result (e.g. Oelmann & Laeng, 2009). Importantly though, we did observe clear group

differences both in MET-scores and in tapping accuracy on both experiments, reducing the

force of the above account.

5.2.2 Musicians outperformed non-musicians in tapping accuracy The tapping results did document musicians’ more efficient processing of polyrhythmic and

microtiming features. In principle, sensorimotor synchronisation is enabled by neural events

that are entrained with the music (e.g., Stupacher et al., 2017). Differences in tapping

accuracy may therefore be seen to reflect differences in neural functionality, engendered by

levels of rhythmic expertise. Across experiments and conditions, the musicians consistently

tapped with substantially higher rhythmic accuracy, despite apparently not requiring more

mental effort than the non-musicians. This clearly illustrate the enhanced rhythm perception

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skills of professional musicians. Interestingly, the relative difference in performance between

groups increased when meter perception ‘challengers’ like polyrhythm and microtiming were

introduced, as we specifically expected: In Experiment 1, despite both groups’ tapping

accuracy decreased from plain to polyrhythm, the negative effect of polyrhythm on tapping

accuracy was more pronounced in the non-musicians than in the musicians. Similarly, in the

microtiming experiment, tapping accuracy was more negatively affected by microtiming in

the non-musicians, although negatively influencing both groups. We may compare our

microtiming tapping results especially to a study by Hove et al. (2007) who found that

occurrence of onset (microtiming) asynchronies increased non-musicians’ tapping variability

(SD of ITI, a negative measure of tapping accuracy), but decreased tapping variability (i.e.

increased accuracy) for the musicians. However, the magnitude of microtiming was larger

(up to 80 ms) in our experiment, than in the Hove et al.-study (up to 50 ms) and only the ±80

ms, and not ±40 ms condition did in fact affect tapping accuracy. In addition, Hove and

colleagues presented two simple sine tones in asynchrony, with an absence of additional

rhythmic cues. Therefore, their musicians might have been able to place the two onsets into a

single internal metric model, aiding their sense of time and increasing their tapping accuracy.

In contrast, there were several rhythmic cues throughout the present study and the bass/drums

asynchrony apparently confused rather than aided beat placement.

5.3 Microtiming and the experience of groove The participants’ response profiles on our two rating scales (‘wanting to move the body to the

music’ and ‘musical well-formedness’) were considerably correlated (r = .56 - .80),

suggesting that an experience of groove (addressed by question 1) is partly an experience of

the groove being musically well-formed (addressed by question 2). Thus, treating the two

scales as one combined score did seem to make both theoretical and empirical sense,

although the two scales may capture unique and subtle aspects of the musical experience.

Also, general ratings of ‘well-formedness’ suggested that participants did not experience the

excerpts as particularly unnatural in musical terms, indicating an adequate ecological validity

of The Microtiming Experiment.

As mentioned earlier, although microtiming is assumed to play a vital role in much

groove-based music, systematic experimental investigations have given inconsistent evidence

as to whether microtiming asynchronies in grooves actually promotes groove experience

(Davies et al., 2013; Kilchenmann & Senn, 2015). Nevertheless, the present findings seem

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rather consistent with earlier studies. Indeed, non-deviated grooves were generally rated

highest on both scales. The larger the microtiming asynchrony magnitudes, the lower the

clips were rated. There could be at least four reasons why the most highly rated grooves were

the on-the-grid versions. First, it might be that systematic microtiming asynchrony between

bass and drums is not a correlate to a groove experience, at least not for the type of grooves

chosen for this experiment. Second, it might be that microtiming in a rhythm section dyad

only fully reaches its potential when the musicians (or producer) intuitively try to groove.

That is, ‘moving around’ mechanically on bass tracks in ProTools is a rather non-musical

way of achieving ‘tasteful’ rhythmic nuances. Roholt (2014) suggests that the analytical

approach of measuring (and in this case manipulating) timing nuances may give little sense

as a way to understand microtiming’s role in groove-based music, as long as measurements

are not connected to a phenomenological account of the groove (i.e., the effect it has on the

listener). An interesting finding from the present study may illustrate this point: the second

highest rated clip was the +40 ms version of the high complexity groove (as shown in Figure

11; the only clip that was rated higher, was the on-the-grid version of the same groove type).

The high complexity groove was the only excerpt based on a real tune, and the +40 ms

condition was the excerpt that resembles most precisely the timing of the bass and drum track

from the original version of Really Love. One might therefore speculate that this particular

excerpt reflects a pattern between bass and drums where microtiming is indeed suitable, and

that the ‘groovy’ feel from the original album was partly re-created in the corresponding

condition of the present experiment. Third, since Roholt (2014) argues that the potential

groove promoting effects of microtiming depend on its role in the musical context and that

music is perceived as a figure/ground kind of relationship, microtiming should not be given

the figure role; instead, to contribute to a groovy experience, it must fade into the

background. The rather repetitive and non-ecological experimental setting could have

contributed to both an analytical rather than a more ‘holistic’ listening mode, with an

enhanced focus on the asynchronies. Fourth, when only two instruments were presented in

asynchrony, this created a confusing sense of two separate, phase-shifted, rhythmic streams.

If the timing of either drums or bass had been supported by additional instruments, like a

synth or a guitar, this would have given more acoustic salience to one of the timings, and

therefore a stronger cue in the listener to experience that timing as the main one, with

asynchrony timing either pushing or pulling against. Said differently, reducing the ambiguity

concerning which time is experienced as the reference time may be a key to experience

microtiming asynchrony as groove promoting.

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Group differences emerged also on subjective ratings. Consistent with earlier studies

(Davies et al., 2013; Senn et al., 2016), musicians showed an increased responsivity to

different microtiming conditions, compared to the non-musicians, with the use of wider

ranges in the rating scales. It is not unknown that musicians outperform non-musicians in

discriminating between fine-scaled rhythmic nuances. For example, while trained listeners

need around 20 ms asynchrony between two tones (in isolation) to decide which of them is

presented first (Hirsh, 1959), untrained listeners usually need longer time intervals

(Broadbent & Ladefoged, 1959). Even if it was plausible to speculate that musical

competence would increase not only responsivity, but also tolerance or even preference for

microtiming (as for example suggested by Hove et al., 2007), such tolerance was only

reflected in tapping data, not in subjective rating or pupil data.

Finally, although the on-the-grid versions were generally the most preferred, we made

the interesting observation that participants seemed to prefer the microtiming conditions

where the double bass was timed after the drums (similar to the original of D’Angelo’s

Really Love), in favour of ahead of the drums. Moreover, -80 ms condition (bass 80 ms ahead

of drums) yielded both the highest effort, lowest tapping accuracy and lowest rating of all

excerpts. It is not clear however, if this finding can be generalized to other musical genres,

such as a bass/drums swing jazz accompaniment.

5.4 Groove experience, effort and complexity A question of interest concerned the processing of microtiming, and involved exploring

possible systematic relations between effort and groove experience, moderated by rhythmic

structural features/complexities. Unravelling such associations is intriguing, but a

considerable endeavour, given the somewhat paradoxical ‘effortless-but-complex’-dialectic

of groove and groove experience. On one hand, music theorists, musicians and listeners may

characterize groove as a seemingly smooth and effortless musical attendance (Danielsen,

2006; Janata et al., 2012; Roholt, 2014), being paralleled with the well-researched concept

(both in music and elsewhere) of flow (Csikszentmihalyi, 1990; de Manzano, Theorell,

Harmat, & Ullén, 2010). On the other hand, groove-based music also admittedly contains

structural complexities that challenge or even violate listeners’ sense of meter (Vuust et al.,

2014; Witek, 2017). Thus, one possibility is that different structural rhythmic features pave

the way for different connections between effort and ratings of groove; allowing results both

supporting the effortless- and the complexity-side of groove.

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Janata et al., (2012) suggest that the affectively positive psychological state of feeling

in the groove is closely linked to perceptual fluency, as individuals are able to anticipate

meter and distinct musical onsets, and actualize or imagine them in relation to body

movements. The authors found participants’ tapping accuracy to be higher when engaging

with music rated as high-groove, compared to low-groove; participants also rated tapping and

movement synchronisation tasks as easier when they at the same time rated the music as

groovier. Hence, high predictability and ‘effortless’ attendance seems to be two related key

aspects to groove experience, since predictability comprises perceptual fluency that is again

related to ‘effortlessness’. Indeed, we found that complexity in the form of microtiming

seemed to increase prediction error in a way that concomitantly increased effort and

decreased groove. The on-the-grid excerpts, being rhythmically most predictable, were the

highest rated, having the highest tapping accuracy and at the same time demanding least

cognitive workload from the participants. When bidirectional microtiming asynchrony

increased, this decreased subjective ratings, tapping accuracy, and strengthened the pupil

response; this pattern emerged both when participants listened, and when they tapped their

perceived beat. The microtiming asynchronies’ effect on pupil size yielded a U-shaped curve

(Figure 7), while the microtiming effect on groove rating gave an inverted U-shaped curve

(Figure 10). It is tempting to propose that microtiming asynchronies in the present

experiment, perhaps with the exception of the most ecologically valid +40ms D’Angelo

excerpt, were experienced by the listeners, not as groove promoters, but more as pure

unmusical ‘metric violations’. Manifested as brain response, the prediction error between

rhythm and meter caused by microtiming asynchrony was signalled by the observed

corresponding increase in effort.

Additional evidence from tapping and subjective rating data supporting the effortless-

notion of a groove experience, was the finding that timekeeping increased both tapping

accuracy and groove ratings; this being consistent with Madison et al. (2011). Timekeeping

probably reduced metric ambiguity and increased predictability, independent of whether

microtiming features was present or absent. Probably, the hi-hat eight-notes provided

listeners with more temporal information and simultaneously added extra salience to the

drum kit timing, making it appear more like the ‘main’ timing, in favour of the timing of the

bass or ‘somewhere between the timing of the bass and drum’ (like discussed in the previous

section). However, we did not observe any significant effect of pupil responses by adding hi-

hat eighth notes, as it could possibly be expected.

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Some of our results also indicate that groove experience does not only emerge from

predictability, but also from complexity. To illustrate: even if a series of metronome click

have extremely high rhythmic predictability and is easy to bodily synchronise with, that does

surely not alone promote groove (Stupacher et al., 2013). It has recently been both

theoretically proposed (Witek, 2017) and empirically shown (Witek et al., 2014) that

structural complexities producing tension between the rhythm and the underlying meter

(without breaking down the listeners’ metrical model), may be groove experience

components by virtue of inviting the participant to participate. This would allow the

‘fulfilling’ with their bodies the open spaces or metrical ambiguities in the groove. Witek et

al. (2014) found an inverted U-shaped curve between syncopation density in drum kit

grooves, and groove experience. Thus, too little and too much syncopation (complexity)

yielded lower groove experience, while medium syncopation was related to higher groove

experience. The findings were thought to reflect a general inverted U-shaped relation

between structural complexity and aesthetic pleasure of art (Berlyne, 1971). Interestingly, in

the present study, higher levels of complexity (numbers of syncopations and sound event per

bar) increased groove ratings, and in addition respondents were more responsive to

microtiming when complexity increased (Figure 11), giving a ‘possibility’ for microtiming to

play a more adaptive role in more complex groove matrices. However, we did not observe a

U-shaped pattern between complexity and groove experience, possibly because grooves with

an ‘exaggerated’ density of syncopations/notes in our stimuli material were not included, as

in Witek and colleagues’ study. This was perhaps an additional reason for finding this

variable to be unrelated to mental effort. A complexity effect emerging from

syncopations/sound events per bar could nevertheless be seen in participants’ tendency to tap

with lower tapping accuracy, even if groove ratings were positively related to complexity.

This last finding contrasts the earlier results by Janata et al., (2012) and underlines the

impression that predictability alone does not necessarily increase groove.

To conclude, different types of rhythmic features (microtiming, ‘meter-supporting’

timekeeping and syncopations) might have yielded different kinds of effects on mental effort,

tapping and subjective ratings – together supporting the ‘effortless-but-complex’-dialectic.

When moderated by microtiming, groove ratings were negatively related to effort, probably

since microtiming was experienced more like a metric violation than a groove promoter. Hi-

hat eighth notes increased predictability and concomitantly yielded higher groove ratings and

tapping accuracy. Yet, under influence of timekeeping, groove ratings were empirically

unrelated to mental effort, since adding a timekeeper did not influence pupil response.

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Syncopations (and note onsets per bar) represented a structural complexity that promoted

groove ratings despite lowering tapping accuracy, but interestingly it did not affect mental

effort. Therefore, in the present experiment, mental effort in musical grooves signal metric

violations that have concomitantly a decreasing effect on groove. It remains an open question

whether the fact that structural features which increased groove ratings were not reflected in

pupil size, can be generalized to other groove-contexts.

5.5 Limitations and methodological considerations As in all scientific investigations, the present study has several limitations, of which some

have been discussed throughout the text. Not yet discussed are possible problems related to

recruitment of participants. Participants were recruited from a personal network of musicians

and non-musicians. Musicians were told that they were invited to participate because they

were musicians, giving them an incentive to ‘perform well’. In contrast non-musicians, who

were selected on the basis of being ‘not musicians’, by being deprived of a certain quality.

Moreover, the experiment was not blinded. The author was the experimenter whom is also a

professional musician. His presence during the experiment could have yielded more

performance anxiety to the musicians.

An obvious limitation of the present study lies in the ecological aspect of the

microtiming stimuli, being all ‘artificially’ designed. This made experimental control

possible, but had several drawbacks related to our research questions (e.g. of microtiming’s

relation to groove). In addition, the ‘Complexity’ variable was confounding syncopation

density with note onsets per bar. We chose this solution, since the processing effects from

syncopations were not the main aim of the present study. There was also a potential source of

bias in that the high complexity groove constituted a copy of an original tune (Really Love),

which several participants perhaps recognized; this was not asked for. Another issue was that

we did not control statistically for musical genre preference. We mapped this in the

questionnaire, but did not see the direct advantage of including it in the study, since the

grooves (at least the ‘low’ and ‘medium’) were quite ‘generic’. Moreover, we did not in

particular address the role of affect and aesthetic experiences in this study, although

discussing it briefly in part 5.2.1. This is nevertheless a limitation, since affect is an important

part of music perception in general and of the experience of groove in particular.

There were technical problems related to tapping data, already discussed in the

‘Methods’-section. The absolute offset tapping data seemed unreliable, so we did not get the

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chance to investigate how microtiming influenced absolute offsets. Clearly, it was non-

optimal to tap on a standard PC-keyboard; a device specially made for tapping would perhaps

have made a significant difference. As for the groove ratings, we might have measured

participants’ actual movements (cf. Kilchenmann & Senn 2015), instead of just asking about

‘wanting to move’. Though, an obvious challenge was that each participant’s head was kept

in a fixed position while listening to music, in order to get optimal pupil data. As was

discussed earlier, although presenting two theories of meter perception (PC and DAT), we do

not have any foundation to say which of the two theories that explain our data better.

5.6 Future studies The present study may spark the interest in several future investigations on the following:

• Syncopations and effort, for example with syncopation stimuli from the study by Witek et

al. (2014), to further investigate the relation between rhythmic complexities, effort and

groove experience, also when syncopation density is ‘exaggerated’.

• The effect of microtiming on effort and ratings in instances where ecological validity and

‘aesthetics’ are maximized, for example by using original tracks.

• The relevant musical contexts for microtiming, that is, the characteristics of musical and

rhythmic patterns that are suitable for microtiming to play a groove-promoting role.

• Replications of the present microtiming study with swung eight-note jazz

accompaniments (e.g. cymbal and bass; see Collier & Collier 1996), instead of the even

eighth grooves of the present study, also to assess generalizations from the present study

to other genres.

• The effect of effort related to free form rather than isochronous tapping. In a free-form-

paradigm, participants could be asked to tap whatever rhythmic pattern to the music that

felt comfortable. The hypothesis is that this should be more effort demanding (Janata et

al., 2012).

• General and expertise effects in effort during silent periods following rhythmic material,

to measure effects of top-down processes only, as done by Stupacher et al. (2017), and

discussed in section 5.2.1.

• Effects from maximizing group differences, for example by comparing professional

drummers/percussionists with participants with dysrhythmia (Launay, Grube, & Stewart,

2014), having specific impairments in meter and beat perception.

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6 Conclusion By addressing relations between local auditory events (rhythm) and the underlying mental

structure (meter) in groove-based music, the present study taps into cognitive and brain

functioning in terms of time perception, anticipation, bottom up/top-down processing,

musical effort and the experience of groove. Further, it adds to the scientific literature of

documenting domain-specific expertise effects of music perception. To our knowledge, it is

the first time an index of mental effort – measured by pupillary response – is employed

during exposure to instances of polyrhythm and microtiming in a groove context. Both

presence of polyrhythm/cross-rhythms and magnitudes of microtiming asynchronies between

bass and drums were found to be positively related to mental effort and negatively related to

tapping accuracy. The main results apparently support Dynamic Attending Theory (DAT)

and Predictive Coding Theory (PC), which both predict that rhythmic instances (complexity)

lowering prediction accuracy, should increase effort and decrease tapping accuracy. Group

differences in pupil response were not observed and possible reasons for this were discussed.

Nevertheless, the perceptual skills related to rhythm perception in professional jazz musicians

were strongly reflected in their superior tapping accuracy, despite not allocating more effort

to the task than the non-musicians. In both experiments, tapping the beat while listening

yielded higher psychophysiological effects (more mental effort) than when listening only.

When giving their subjective ratings of groove and well-formedness, participants

generally preferred the on-the-grid grooves more than the microtiming grooves, consistent

with existing research. However, grooves that reflected a rhythm matrix in which

microtiming had been suitable in a real musical setting did yield high ratings also in the

corresponding microtiming condition. Timing conditions where bass was timed behind drums

were more preferred than conditions with bass timed ahead of drums. Musicians were more

responsive (used a bigger ranger of the subjective rating scale) but less tolerant to

microtiming, than the non-musicians. Finally, we proposed that the link between mental

effort and groove experience, moderated by rhythmic complexity, is dependent on ‘type’ of

complexity. In our stimuli material, rhythmic features that increased groove were not

reflected in effort, even if structural complexity (syncopations) was added. In contrast, the

rhythmic complexity of microtiming asynchronies decreased groove and was related to

higher effort, possibly functioning more like a pure metrical violation than a groove

promoter.

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Wu, E. X. W., Laeng, B., & Magnussen, S. (2012). Through the eyes of the own-race bias:

Eye-tracking and pupillometry during face recognition. Social Neuroscience, 7(1–2),

202–216. http://doi.org/10.1080/17470919.2011.596946

Zatorre, R. J., Chen, J. L., & Penhune, V. B. (2007). When the brain plays music: auditory-

motor interactions in music perception and production. Nature Reviews. Neuroscience,

8(7), 547–558. http://doi.org/10.1038/nrn2152

7.2 Discography D’Angelo (2000). Voodoo. RCA.

Destiny’s Child (2004). Survivor. Columbia.

Jack Ü, Skrillex, Diplo (2015). Skrillex and Diplo presents Jack Ü. WEA International.

Keita, Salif (1987). Soro. Island Records.

Spears, Britney (2011). Femme Fatale. JIVE records.

Sting (1987). Nothing Like the Sun. A&M Records.

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7.3 Filmography

Collins, A. (2014). How playing an instrument benefits your brain. TED Education.

Retrieved from http://ed.ted.com/lessons/how-playing-an-instrument-benefits-your-

brain-anita-collins

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8 Appendix 8.1 Figures A1-A7

Figure A1. Experiment 1: Overview of ‘the polyrhythmic condition’; an excerpt from Sting’s Lazarus Heart (1987). Adapted from Vuust et al., (2006).

Figure A2. Experiment 2: Low structural complexity groove without hi-hat (‘low’)

Figure A3. Experiment 2: Low structural complexity groove with hi-hat (‘low HH’). See Figure A2 for further details on tempo, timing and instrumentation.

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Figure A4. Experiment 2: Medium structural complexity groove without hi-hat (‘medium’). See Figure A2 for further details on tempo, timing and instrumentation.

Figure A5. Experiment 2: ‘Medium structural complexity groove with hi-hat (‘medium HH’). See Figure A2 for details on tempo, timing and instrumentation.

Figure A6. Experiment 2: High structural complexity groove (‘high HH’). See Figure A2 for details on tempo, timing and instrumentation.

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Figure A7. Stereo waveform (amplitude/time) illustrating the microtiming asynchrony between bass and drums in D’Angelo’s Really Love (2014). The onsets of the kick drum and the bass are marked. Made by AD.

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8.2 Tables A1-A5 Table A1 Overview of experimental procedure

Note: Order of stimuli was counterbalancd for half of the participants in each group (musicians/non-musicians).

Table A2 Tapping exclusions in Experiment 1

Note: Tapping exclusions were calculated according to rules accounted for in the ‘Data processing’-section. Table A3 Tapping exclusions in Experiment 2 (overview from Table A4 & A5)

Note: Tapping exclusions were calculated according to rules accounted for in the ‘Data processing’-section.

Overview of experimental procedureExperiment Condition Trial

Experiment 2: Microtiming 'Listen only', rate excerpts after listening 1-25

Experiment 1: Polyrhythm vs. non-polyrhythm 'Listen only', rate excerpts after listening 1-2

BREAK

Experiment 1: Polyrhythm vs. non-polyrhythm Tap with the beat 25-1

Experiment 2: Microtiming Tap with the beat 2-1

BREAK

Musical Ear Test 1-52

Tapping exclusions in Experiment 1’the polyrhythmic condition’

’the plain condition’

Total

Musicians

Excluded trials 1 0 1

Missing reference points in included trials 3 0 3

Double taps in included trials 5 3 8

Non-musicians

Excluded trials 4 2 6

Missing reference points in included trials 10 4 14

Double taps in included trials 14 10 24

Tapping exclusions in Experiment 2 (overview from Table A4 & A5)Musicians Non-musicians

Excluded trials 5 63

Missing reference points in included trials 149 109

Double taps in included trials 32 296

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Table A4 Tapping exclusions in Experiment 2, across ‘Microtiming’ and ‘Complexity’-conditions

Note: Tapping exclusions were calculated according to rules accounted for in the ‘Data processing’-section.

Tapping exclusions in Experiment 2, across ‘Microtiming’ and ‘Complexity’-conditionsMusicians ‘low’ ‘low HH’ ‘medium’ ‘medium HH’ ‘high HH’ Total-80 ms

Excluded trials 2 0 0 0 0 2

Missing reference points in included trials 2 4 13 1 16 36

Double taps in included trials 2 0 1 3 0 6

-40 ms

Excluded trials 0 0 1 0 0 1

Missing reference points in included trials 4 3 6 5 13 31

Double taps in included trials 1 2 0 2 2 7

0 ms

Excluded trials 0 0 0 0 0 0

Missing reference points in included trials 3 2 4 4 14 27

Double taps in included trials 2 0 0 1 0 3

+40 ms

Excluded trials 0 0 0 0 0 0

Missing reference points in included trials 7 1 4 2 15 29

Double taps in included trials 5 5 2 0 0 12

+80 ms

Excluded trials 1 0 0 0 1 2

Missing reference points in included trials 4 2 2 1 17 26

Double taps in included trials 2 1 0 1 0 4

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Table A5 Tapping exclusions in Experiment 2, across ‘Microtiming’ and ‘Complexity’-conditions

Note: Tapping exclusions were calculated according to rules accounted for in the ‘Data processing’-section.

Tapping exclusions in Experiment 2, across ‘Microtiming’ and ‘Complexity’-conditionsTotal Non-musicians ‘low’ ‘low HH’ ‘medium’ ‘medium HH’ ‘high HH’ Total

-80 ms

2 Excluded trials 5 4 5 2 1 17

36 Missing reference points in included trials 2 2 11 4 10 29

6 Double taps in included trials 9 15 15 18 21 78

-40 ms

1 Excluded trials 2 6 3 1 1 13

31 Missing reference points in included trials 4 0 3 2 5 14

7 Double taps in included trials 14 5 14 1 2 36

0 ms

0 Excluded trials 2 2 2 3 3 12

27 Missing reference points in included trials 3 1 3 5 11 23

3 Double taps in included trials 19 7 4 8 21 59

+40ms

0 Excluded trials 3 1 0 2 3 9

29 Missing reference points in included trials 1 2 7 0 7 17

12 Double taps in included trials 7 9 15 6 20 57

+80ms

2 Excluded trials 3 0 2 3 4 12

26 Missing reference points in included trials 4 4 4 2 12 26

4 Double taps in included trials 17 9 13 12 15 66