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Neural Entrainment to the Rhythmic Structure of Music
Adam Tierney and Nina Kraus
Abstract
■The neural resonance theory of musical meter explains
musical beat tracking as the result of entrainment of neural
oscillations to the beat frequency and its higher harmonics. This
theory has gained empirical support from experiments using
simple, abstract stimuli. However, to date there has been no
empirical evidence for a role of neural entrainment in the
perception of the beat of ecologically valid music. Here we
presented participants with a single pop song with a super-
imposed bassoon sound. This stimulus was either lined up with
the beat of the music or shifted away from the beat by 25% of
the average interbeat interval. Both conditions elicited a neural
response at the beat frequency. However, although the on-the-
beat condition elicited a clear response at the first harmonic of
the beat, this frequency was absent in the neural response to
the off-the-beat condition. These results support a role for neural
entrainment in tracking the metrical structure of real music and
show that neural meter tracking can be disrupted by the pre-
sentation of contradictory rhythmic cues. ■
INTRODUCTION
Temporal patterns in music are organized metrically, with
stronger and weaker beats alternating. This alternation
takes place on multiple timescales, resulting in a complex
sequence of stronger and weaker notes. Position within
the metrical hierarchy affects how listeners perceive
sounds; strong metrical positions are associated with
higher goodness-of-fit judgments and enhanced duration
discrimination (Palmer & Krumhansl, 1990). The musical
beat is perceived where strong positions at multiple time-
scales coincide, although individual differences exist in
the scale at which listeners perceive the beat (Iversen
& Patel, 2008; Drake, Jones, & Baruch, 2000).
Metrical processing begins early in life: Brain responses
to rhythmic sounds in newborn infants are modulated by
each soundʼs position in the metrical hierarchy (Winkler,
Haden, Ladinig, Sziller, & Honing, 2009). Metrical per-
ception is, therefore, a fundamental musical skill, and as
such there have been numerous attempts to model how
listeners track metrical structure. An influential model
proposes a bank of neural oscillators entraining to the
beat (Velasco & Large, 2011; Large, 2000, 2008; Van Noorden
& Moelants, 1999; Large & Kolen, 1994), resulting in saliency
oscillating on multiple timescales (Barnes & Jones, 2000;
Large & Jones, 1999). This model is supported by work
showing that beta oscillations are modulated at the rate
of presentation of rhythmic stimuli (Fujioka, Trainor,Large,
& Ross, 2012), possibly reflecting auditory–motor cou-
pling, as well as work showing enhanced perceptual dis-
crimination and detection when stimuli are aligned with
a perceived beat (Bolger, Trost, & Schön, 2013; Miller,
Carlson, & McAuley, 2013; Escoffier, Sheng, & Schirmer,
2010; McAuley & Jones, 2003; Jones, Moynihan, MacKenzie,
& Puente, 2002; Barnes & Jones, 2000).
There is, however, no direct evidence for neural en-
trainment to metrical structure in real music. (We define
“neural entrainment”in this paper as phase-locking of
neural oscillations to the rhythmic structure of music.)
Most investigations of the neural correlates of rhythm pro-
cessing have used simple stimuli such as tone sequences
and compared evoked responses to stimuli in strong and
weak metrical positions. Studies of simple stimuli have
found that strong metrical percepts are associated with
larger evoked potentials and higher-amplitude evoked
and induced beta and gamma oscillations (Schaefer, Vlek,
& Desain, 2011; Vlek, Gielen, Farquhar, & Desain, 2011;
Fujioka, Zendel, & Ross, 2010; Geiser, Sandmann, Jäncke,
& Meyer, 2010; Abecasis, Brochard, del Río, Dufour, &
Ortiz, 2009; Iversen, Repp, & Patel, 2009; Ladinig, Honing,
Háden, & Winkler, 2009; Potter, Fenwick, Abecasis, &
Brochard, 2009; Winkler et al., 2009; Pablos Martin et al.,
2007; Abecasis, Brochard, Granot, & Drake, 2005; Snyder
& Large, 2005; Brochard, Abecasis, Potter, Ragot, & Drake,
2003). Studies of simple stimuli have also demonstrated
neural entrainment to a perceived beat and its harmonics
(Nozaradan, Peretz, & Mouraux, 2012; Nozaradan, Peretz,
Missal, & Mouraux, 2011). Furthermore, a recent study
has shown that alignment with the beat of real, ecologically
valid music modulates evoked responses to a stimulus
(Tierney & Kraus, 2013a) such that on-the-beat stimuli
elicit larger P1 responses; however, this result can either
be attributed to enhancement of processing of the target
stimulus or to neural tracking of the beat of the music.
Thus, no study to date has demonstrated neural entrain-
ment to the rhythmic structure of real music.
Northwestern University
© 2014 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 27:2, pp. 400–408
doi:10.1162/jocn_a_00704
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