Neuroscience 250 (2013) 49–59
THE EFFECTS OF MUSIC ON BRAIN FUNCTIONAL NETWORKS: A
NETWORK ANALYSIS
J. WU, a,b J. ZHANG, a,b X. DING, c R. LI a,b AND
C. ZHOU a,b*
a
Cognitive Science Department, Xiamen University, Xiamen, China
b
Fujian Key Laboratory of the Brain-like Intelligent Systems,
Xiamen University, Xiamen, China
c
College of Foreign Languages and Cultures, Xiamen
University, Xiamen, China
Abstract—The human brain can dynamically adapt to the
changing surroundings. To explore this issue, we adopted
graph theoretical tools to examine changes in electroen-
cephalography (EEG) functional networks while listening
to music. Three different excerpts of Chinese Guqin music
were played to 16 non-musician subjects. For the main fre-
quency intervals, synchronizations between all pair-wise
combinations of EEG electrodes were evaluated with phase
lag index (PLI). Then, weighted connectivity networks were
created and their organizations were characterized in terms
of an average clustering coefficient and characteristic path
length. We found an enhanced synchronization level in the
alpha2 band during music listening. Music perception
showed a decrease of both normalized clustering coefficient
and path length in the alpha2 band. Moreover, differences in
network measures were not observed between musical
excerpts. These experimental results demonstrate an
increase of functional connectivity as well as a more random
network structure in the alpha2 band during music percep-
tion. The present study offers support for the effects of
music on human brain functional networks with a trend
toward a more efficient but less economical architecture.
Ó 2013 IBRO. Published by Elsevier Ltd. All rights reserved.
Key words: electroencephalography, music, graph theory,
small-world network, synchronization.
INTRODUCTION
The brain is considered to be a complex system,
comprised of spatially interconnected areas. Structural
and functional connectivity of the brain possess
properties of complex networks and can be investigated
*Correspondence to: C. Zhou, Cognitive Science Department, Xia-
men University, 422 South Siming Road, Xiamen 361005, China. Tel:
+86-13850083875; fax: +86-592-2580168.
E-mail address: dozero@xmu.edu.cn (C. Zhou).
Abbreviations: DSI, diffusion spectrum imaging; DTI, diffusion tensor
imaging; EEG, electroencephalography; EOG, electrooculogram; FIR,
finite impulse response; fMRI, functional magnetic resonance imaging;
GLS, GuangLingSan; MEG, magnetoencephalography; MHSN,
MeiHuaSanNong; OLWJ, OuLuWangJi ; PLI, phase lag index; PSD,
power spectral density; SD, standard deviation.
0306-4522/13 $36.00 Ó 2013 IBRO. Published by Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.neuroscience.2013.06.021
49
using graph theoretical measures (Stam and Reijneveld,
2007; Bullmore and Sporns, 2009; He and Evans, 2010;
Bullmore and Bassett, 2011). One effective model is
small-world networks in terms of graph theory. Small-
world networks appear as regular, lattice-like structures
with a few random long-range connections (Watts and
Strogatz, 1998). They are characterized by high local
clustering, signifying dense local connectivity, and short
path length between any two vertices. Small-world
organizations are interesting for complex brain networks
since they confer high local and global efficiency of
information transfer with a low wiring cost (Bullmore and
Sporns, 2009; He and Evans, 2010).
There is increasing evidence for small-world
configurations in structural networks of animals (Sporns
and Zwi, 2004; Sporns et al., 2007). These
configurations are extended to studies of human brain
structural networks based on cortical thickness
measurements (He et al., 2007), diffusion tensor imaging
(DTI) (Iturria-Medina et al., 2008; Yan et al., 2011) and
diffusion spectrum imaging (DSI) (Hagmann et al.,
2008). In several functional magnetic resonance imaging
(fMRI) studies, networks of functional connectivity also
exhibited small-world characteristics (He et al., 2009;
Hayasaka and Laurienti, 2010; Tian et al., 2011). While
neuronal activities frequently occur in time-scales shorter
than those reflected in fMRI signal fluctuations (Palva
et al., 2010), the use of electroencephalography (EEG)
and magnetoencephalography (MEG) guarantees an
adequate temporal resolution. And such dynamic
techniques are inherently appropriate to detect
oscillatory synchronization (Sporns et al., 2000), which
takes a crucial role in interregional communication
(Fries, 2005; Schnitzler and Gross, 2005). Network
analyses of EEG and MEG are consistent with the fMRI
data, indicating small-world features in synchronization
patterns (Stam et al., 2007a, 2009; Deuker et al., 2009;
Douw et al., 2011; Jin et al., 2012). The small-world
network attributes could be found not only in resting but
also during task conditions, such as finger tapping and
music listening (Eguı́luz et al., 2005). Interestingly, brain
maturation may be associated with a shift toward a more
small-world network architecture (Boersma et al., 2011).
In contrast, brain networks deviate from the optimal,
normal small-world organization in brain pathology (Stam
et al., 2007a, 2009; Rubinov et al., 2009).
It has been suggested that brain networks exist in a
critical state between chaos and order (Kitzbichler et al.,
2009; Petermann et al., 2009). Small-world properties
support rapid adaptive reconfiguration of functional
50 J. Wu et al. / Neuroscience 250 (2013) 49–59
connectivity in response to varying cognitive demands
(Bassett et al., 2006). Several empirical studies have
identified task-related changes in network organization
during finger movements (Bassett et al., 2006; Jin et al.,
2012), working memory (Palva et al., 2010; Kitzbichler
et al., 2011) and learning (Bassett et al., 2011).
Another potential task that can cause network
reconfiguration is music perception. Music is widely
recognized as a universal language. Music in various
forms has a profound impact on all cultures, and even
on animals (Gray et al., 2001). Listening to music is a
complex cognitive task and implicates processing and
integration of meaningful elements including harmony,
rhythm or melody (Schmithorst, 2005). Modulating
effects of music have been observed on functional
connectivity such as default mode network (DMN) (Kay
et al., 2012) and oscillatory synchronization (Petsche
et al., 1997; Bhattacharya et al., 2001; Bhattacharya
and Petsche, 2005; Flores-Gutiérrez et al., 2007, 2009;
Ruiz et al., 2009). Wu et al. (2012) have identified a
topological change in EEG alpha-band networks in a
music listening task while preserving the small-world
characteristics comparable to the findings of Eguı́luz
et al. (2005).
While the previous study has reported network
reconfiguration during music perception, three issues
demand further attention: (i) the topological differences
between musical excerpts have not been considered; (ii)
the earlier analysis was focused merely on the alpha
band, whereas the functional networks were usually
probed in multiple frequency intervals (Deuker et al.,
2009; Stam et al., 2009; Boersma et al., 2011;
Kitzbichler et al., 2011; Jin et al., 2012) since different
frequency components might represent different
biological significances; (iii) estimation of brain functional
connectivity from EEG recordings can be affected by
volume conduction, which may make interpretation
unreliable.
The problem of volume conduction refers to the
inclination of nearby EEG electrodes to pick up activity
of common neural sources, introducing high synchronies
that do not reflect factual functional connectivity (Nunez
et al., 1997; Stam et al., 2007b). There are two main
approaches proposed to tackle this problem. A first
approach attempts to use reconstructed sources as a
basis to assess functional correlations (Gross et al.,
2001; Lehmann et al., 2006; Amor et al., 2009).
Although this method indeed makes a contribution to
detecting functional co-operations between anatomically
well-specified brain areas, a primary defect is the
absence of a unique solution to the EEG inverse
problem. A second approach is to determine true
interactions between time series which are not likely to
be biased by volume conduction. One could adopt the
imaginary part of the complex coherency, which is not
owing to volume conduction effects (Nolte et al., 2004).
However, it contains mixed information on both
amplitudes and phase delays, and consequently is not
an appropriate measure of functional correlations. Stam
et al. (2007b) proposed phase lag index (PLI) as an
alternative. The PLI estimates consistency of phase
difference between two time series. It can determine a
reliable evaluation of synchronization that is less
sensitive to the influence of volume conduction.
Therefore we utilize the PLI to quantify functional
connectivity.
In this study, we intend to further explore the effects of
music on EEG functional networks. Our questions are: (i)
whether network reorganization takes place during music
perception in different frequency bands and (ii) whether
differences in network organization can be observed
between musical pieces. The experiment used three
pieces of Chinese Guqin music as musical stimuli.
Guqin music is regarded as a symbol of Chinese
civilization (Zhu et al., 2008, 2009) and has been
selected as a Masterpiece of the Oral and Intangible
Heritage of Humanity by UNESCO. Cognitive
neuroscientific research on Guqin music perception will
deepen our understanding of the perception of Eastern
music.
EXPERIMENTAL PROCEDURES
Participants and materials
This study involved 16 right-handed subjects (mean age
22.25 years, standard deviation (SD) 1.65; eight males).
All subjects were recruited from the university campus
and nurtured in China. Structured interviews guaranteed
that the subjects had normal hearing, no formal or
informal training in music, and were free from
neurological diseases or psychoactive drugs use. Each
subject was informed of the experimental procedure and
gave a written consent to participate. The subjects were
paid for the participation in the experiment.
All subjects were instructed to listen attentively to
three pieces of Guqin music: OuLuWangJi (OLWJ),
MeiHuaSanNong (MHSN), and GuangLingSan (GLS)
(music conditions). These selected musical pieces were
unfamiliar to all participants. In addition, subjects also
listened to a segment of pink noise (noise condition).
The presentation of each acoustic stimulus lasted 40 s
and their order was counterbalanced among subjects.
EEG at rest was recorded for 2 min and used as a
control condition (silence condition). The subjects were
comfortably seated inside a dimly illuminated and sound
attenuated chamber. The sound backgrounds were
presented in two stereo speakers located at 2 m in front
of the subjects. The volume arriving at the subjects was
adjusted to 60 dB SPL. Subjects were asked to keep
their eyes closed during the experiment.
EEG recordings
EEG was recorded using the Neuroscan system
(Neuroscan Inc., El Paso, TX) with 64 electrodes
arranged according to the international 10–20 system.
All leads were referenced to left and right earlobes. To
monitor eye movements and blinks, horizontal and
vertical electrooculograms (EOGs) were recorded from
two bipolar electrodes placed slightly lateral to the outer
canthus of each eye and above and below the left eye.
Inter-electrode impedance levels were kept below 5 kX.
 J. Wu et al. / Neuroscience 250 (2013) 49–59
EEG was continuously recorded with a 0.05–100 Hz
bandpass filter, at a 1000 Hz sampling frequency.
Ocular artifacts were reduced by means of a regression
approach (Semlitsch et al., 1986) implemented in Scan
4.3 (Neuroscan Inc., El Paso, TX). In order to avoid
transition effects, the first and last 5-s data of each
music and noise segments were discarded from the
stored EEG. Thus, for each subject, we chose one
artifact-free epoch of 30 s (30,000 samples) for each
task as well as for rest. Using a causal finite impulse
response (FIR) filter (24 dB/octave roll-off), we digitally
filtered the signal in each epoch into the classic
frequency bands: delta (0.1–4 Hz), theta (4–8 Hz),
alpha1 (8–10 Hz), alpha2 (10–13 Hz), beta (13–30 Hz),
and gamma (30–45 Hz).
Power spectral analysis
For each epoch and every electrode, the power spectral
density (PSD) of EEG was obtained via Welch’s method
of spectral estimation (eight equal-length sections with
50% overlap, Hamming window). The PSD values were
averaged across all EEG channels, resulting in a single
value for each band. Power spectral analysis,
calculations of the PLI, network analysis and statistical
analysis were implemented in Matlab (MathWorks Inc.,
Natick, MA).
Calculation of phase lag index
We quantified synchronizations between all pair-wise
combinations of EEG channels via the PLI. The PLI
extracts consistency of phase synchrony between two
signals. It is less sensitive to the contribution from
volume conduction and thus can provide a reliable
appraisal of coupling strength (Stam et al., 2007b;
Pearce et al., 2010). Moreover, the PLI is inherently
suitable to deal with nonlinear and non-stationary
signals like EEG.
For any arbitrary EEG signal x(t), the analytic signal
w(t) is constructed by a complex function of time:
Z þ1
xðsÞ
wðtÞ ¼ xðtÞ þ ixH ðtÞ  xðtÞ þ ip1 p:v: ds; ð1Þ
1 ts
where xH ðtÞ is the Hilbert transform of x(t), and p.v. denotes the
Cauchy principal value. Hilbert transform is virtually the
convolution of x(t) with 1/pt. The instantaneous amplitude A(t)
and instantaneous phase /ðtÞ of x(t) can be derived in terms of
the analytic signal’s polar form:
wðtÞ ¼ AðtÞei/ðtÞ : ð2Þ
In turn, the phase is uniquely determined as
xH ðtÞ
/ðtÞ ¼ arctan : ð3Þ
xðtÞ
From the phases of two EEG signals xa(t) and xb(t),
the phase difference or relative phase is formed as
uab ðtÞ ¼ /a ðtÞ  /b ðtÞ: ð4Þ
Then, the PLI is defined as an asymmetry measure for
the phase difference distribution via
 
1 X
N1  density of the entire graph.
 
PLIab ¼ signðuab ðnÞÞ; ð5Þ
N n¼0 
51
where function signðuab ðnÞÞ returns 1 if uab ðnÞ is positive, and
returns 1 if uab ðnÞ is negative; N denotes the amount of
sampling points. The value of PLIab is restricted between 0 and
1. It reaches 0 for no co-ordination or coupling with a relative
phase that encircled 0 mod p, which is likely to result from
volume conduction. Whereas PLIab = 1 in the condition of strict
phase locking at a constant, nonzero lag.
The PLI values were averaged over all pair-wise
combinations of EEG electrodes to indicate the mean
levels of phase synchronization of cortical areas. In
addition to computing the global mean of PLI, a more
detailed, regional approach was adopted. For this, EEG
electrodes were grouped into frontal, central, parietal,
occipital and temporal areas for both hemispheres.
Averaging was done to obtain intraregional (short-
distance) and interregional (long-distance) PLI.
Intraregional PLI was calculated as the mean value of PLI
for all electrode pairs within one area, and interregional
PLI was calculated between electrodes from two different
areas. Midline channels were not involved.
Network analysis
A network can be typically represented by a graph,
defined as a set of vertices linked by edges. In this
study, we constructed weighted graphs with 62 vertices
(corresponding to the 62 available EEG channels) and
employed the matrices of PLI values for all pairs of
EEG channels as edge weights. That is, we used
62  62 correlation matrices, and the value of each
element wij was assigned as the weight of the edge
between two vertices i and j. The procedure involved in
weighted graph analysis applied to the EEG data is
shown in Fig. 1.
The clustering coefficient CW and the characteristic
path length LW were utilized to quantify the topological
properties of weighted graphs. We adopted weighted
network measures so as to make full use of information
in the weights and avoid the arbitrary selection of
threshold for PLI values. See Stam and Reijneveld
(2007) for an in-depth discussion.
In principle, the clustering coefficient for a vertex i
quantifies the proportion of its neighboring vertices j that
are also connected to each other. Its calculation from
weighted graphs demands weight symmetry (wij = wji)
and 0 6 wij 6 1. These two conditions are satisfied when
employing PLI as edge weights. The clustering
coefficient CW of vertex i can be formalized as (Stam
i
et al., 2009)
P P
j–i k–i wij wik wjk
CW k–j
¼ P P : ð6Þ
i
k–i wij wik
j–i k–j
CW is the mean clustering coefficient over all vertices,
computed as
1X M
CW ¼ CW ; ð7Þ
M i¼1 i
where M denotes the number of vertices. CW depicts the local
We applied the notion of the global efficiency E
(Latora and Marchiori, 2001) to compute the path length
52 J. Wu et al. / Neuroscience 250 (2013) 49–59

Procedure for weighted graph analysis

A B

calculation of
phase lag index

D C FP1-ch.1
0.6 0.6
F2-ch.11
0.5 0.5
FC4-ch.21
0.4 0.4
C6-ch.31
0.3 0.3
TP8-ch.41
0.2 0.2
PO7-ch.51
0.1 0.1

O2-ch.61 0
0
ch.1 ch.11 ch.21 ch.31 ch.41 ch.51 ch.61

randomization
CW LW

E 0.6

0.5
F γ=CW/CW-s C -s
W
0.4

λ=LW/LW-s 0.3
LW-s
0.2

0.1

Fig. 1. Schematic representation of the procedure involved in weighted graph analysis of EEG recordings. EEG signals are recorded from
electrodes illustrated in (A). Epochs of EEG data are filtered within frequency bands of interest (B). PLI is estimated as a measure of correlation
between all pairs of channels, resulting in correlation matrices (C). Then, weighted graphs are constructed based on the correlation matrices (D).
The clustering coefficient CW and characteristic path length LW are calculated to characterize each graph. In addition, surrogate graphs are derived
by randomly shuffling the cells of the correlation matrices (E). For each original graph, network parameters are assessed and averaged over the
ensemble of surrogate graphs, resulting in CW-s and LW-s. Finally, the normalized parameters CW/CW-s and LW/LW-s are determined (F).

of weighted graphs. The length of an edge is determined
as the reciprocal of its edge weight, i.e., Lij = 1/wij if
wij – 0, and Lij = 1 if wij = 0. The distance dij between
the vertices i and j is the minimum of the sum of edge
lengths between these two vertices. The characteristic
path length LW of the whole graph is the average dij for
all possible pairs of vertices. This definition employs the
harmonic mean to handle the problem of disconnected
graphs (Newman, 2003), for which dij ? 1. That is,
1 1
LW ¼ ¼ P PM 1 :
E ð1=MðM  1ÞÞ Mi¼1 j¼1 d
ij
j–i
W
L reflects the overall integration of the graph.
Each network has different basic characteristics, such
as structure, size, and edge weights, influencing both
values of CW and LW. In order to derive network
parameters that are independent of differences in the
PLI, CW and LW were compared to the average values
for an ensemble of surrogate graphs. Fifty surrogate
graphs were constructed from each original graph by
randomly shuffling the edge weights. The normalized
parameters were calculated as
c ¼ CW =CW -s and k ¼ LW =LW -s; ð9Þ
ð8Þ
where CW-s and LW-s denoted the mean CW and LW over 50
surrogate random graphs.
 J. Wu et al. / Neuroscience 250 (2013) 49–59 53

Table 1. Average PSD while exposure to five sound backgrounds in six frequency bands

Bands OLWJ MHSN GLS Noise Silence

Mean ± SD Mean ± SD Mean ± SD Mean ± SD Mean ± SD

Delta 40.6767 ± 24.4940 46.0428 ± 24.0124 39.8405 ± 35.8843 60.9527 ± 43.5244 64.5141 ± 43.1975
Theta 1.4757 ± 0.8573 1.3639 ± 0.9402 1.4023 ± 0.9736 1.6919 ± 0.9183 2.3405 ± 1.8314
Alpha1 2.5426 ± 2.0330 2.4717 ± 1.9757 2.5167 ± 2.0821 2.2861 ± 1.8435 2.5738 ± 1.7728
Alpha2 1.1471 ± 0.4437 1.0981 ± 0.5016 0.9555 ± 0.3833 0.9754 ± 0.3794 1.0923 ± 0.4162
Beta 0.2640 ± 0.0740 0.2582 ± 0.0790 0.2598 ± 0.0777 0.3473 ± 0.1296 0.4207 ± 0.2848
Gamma 0.0253 ± 0.0106 0.0233 ± 0.0081 0.0247 ± 0.0077 0.0393 ± 0.0127 0.0422 ± 0.0313

Weighted EEG networks

OLWJ MHSN GLS Noise Silence
0.3
0.25
0.2
Delta

0.15
0.1
0.05
0
0.25
0.2
Theta

0.15
0.1
0.05
0
0.4
0.35
0.3
Alpha1

0.25
0.2
0.15
0.1
0.05
0
0.4
0.35
0.3
Alpha2

0.25
0.2
0.15
0.1
0.05
0
0.25
0.2
Beta

0.15
0.1
0.05
0
0.1
0.09
0.08
Gamma

0.07
0.06
0.05
0.04
0.03
0.02
0.01
0

Fig. 2. Mean weighted networks in six frequency bands for five sound backgrounds: OLWJ, MHSN, GLS, noise, and silence. The strength of PLI for
all pairs of EEG channels is indicated with a jet scale, from blue (PLI = 0) to red (high PLI). (For interpretation of the references to color in this figure
legend, the reader is referred to the web version of this article.)

Statistical analysis comparisons was applied (i.e., P values < 0.05/

Since parametric statistical analyses might not be
applicable for the subject size, non-parametric methods
were carried out. The PSD, PLI and network measures
among background conditions were compared by
Wilcoxon signed rank test. As the possible 10
correlations among five sound backgrounds
(C25 ¼ 5  4=2 ¼ 10) were subjected to multiple non-
independent tests, Bonferroni correction for multiple
10 = 0.005 were considered as significant).
RESULTS
Statistical comparisons for average PSD during exposure
to five sound backgrounds were performed with the non-
parametric Wilcoxon signed rank test in six different
frequency bands. The results are shown in Table 1. No
54 J. Wu et al. / Neuroscience 250 (2013) 49–59

significant differences were detected among the sound
backgrounds.
Fig. 2 shows the mean graphs for different
background conditions calculated with phase lag index
in six frequency bands. Sketchy differences between the
music and noise conditions and between the music and
silence conditions could be observed via visual scrutiny,
particularly in the alpha1 and alpha2 bands.
The mean PLI values averaged across all pairs of
EEG channels are shown in Table 2. There were
significant differences mainly in the alpha2 band. The
synchronization levels for alpha2 band were significantly
higher during listening to MeiHuaSanNong (P = 0.0019)
and GuangLingSan (P = 0.0032) excerpts in relation to
the noise condition. Increases within alpha2 frequency
were found during listening to OuLuWangJi
(P = 0.0045), MeiHuaSanNong (P = 0.0045) and
GuangLingSan (P = 0.0023) as compared to the
silence condition. Although OLWJ showed a higher
value in comparison with the noise condition
(P = 0.0052) in the alpha2 band, this difference did not
reach significance after Bonferroni correction for multiple
comparisons (a = 0.05/10 = 0.005). In addition, similar
non-significant trends were observed in the alpha1
band. The mean PLI values were higher in the music
conditions than in the noise condition: for OLWJ

Table 2. Mean PLI during exposure to five sound backgrounds in six frequency bands

Bands OLWJ MHSN GLS Noise Silence

Mean ± SD Mean ± SD Mean ± SD Mean ± SD Mean ± SD

Delta 0.1281 ± 0.0188 0.1340 ± 0.0162 0.1386 ± 0.0269 0.1520 ± 0.0260 0.1548 ± 0.0196
Theta 0.0983 ± 0.0334 0.1267 ± 0.0486 0.1149 ± 0.0454 0.0954 ± 0.0265 0.0882 ± 0.0241
Alpha1 0.2245 ± 0.0738 0.2380 ± 0.0862 0.2241 ± 0.0801 0.1632 ± 0.0485 0.1497 ± 0.0434
Alpha2 0.2328b ± 0.0702 0.2452a,b ± 0.0834 0.2300a,b ± 0.0779 0.1566a ± 0.0483 0.1499b ± 0.0406
Beta 0.0986 ± 0.0322 0.1118 ± 0.0401 0.1078 ± 0.0364 0.0767 ± 0.0233 0.0733 ± 0.0206
Gamma 0.0354 ± 0.0048 0.0363 ± 0.0054 0.0340 ± 0.0046 0.0338 ± 0.0042 0.0331 ± 0.0051
For each band, letter superscripts indicate statistically significant differences between backgrounds (Wilcoxon signed rank test, P < 0.005, Bonferroni-corrected).
a
Significant differences between music and noise conditions.
b
Significant differences between music and silence conditions.

Regional differences in phase lag index

Alpha2
OLWJ vs. Noise MHSN vs. Noise GLS vs. Noise

F F F F F F

T C C T T C C T T C C T

P P P P P P

O O O O O O

OLWJ vs. Silence MHSN vs. Silence GLS vs. Silence

F F F F F F

T C C T T C C T T C C T

P P P P P P

O O O O O O

Fig. 3. Robust differences in interregional (denoted by solid lines) and intraregional (denoted by filled squares) PLI for the alpha2 band (Wilcoxon
signed rank test, P < 0.005). The statistical results can only be considered as indicative findings. Solid lines or filled squares in red: sound
backgrounds in red higher than backgrounds in blue; solid lines or filled squares in blue: backgrounds in blue higher than those in red. In comparison
with the noise and silence conditions, the music conditions displayed increased fronto-parietal, fronto-temporal, centro-parietal, centro-occipital and
temporo-parieto-occipital PLI. Intraregional synchronizations for frontal, central and temporal areas were also higher in the music conditions. (For
interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)
 J. Wu et al. / Neuroscience 250 (2013) 49–59 55

Network differences (alpha2 band)

P8
OLWJ vs. Noise MHSN vs. Noise GLS vs. Noise OLWJ vs. MHSN OLWJ vs. GLS

OLWJ vs. Silence MHSN vs. Silence GLS vs. Silence MHSN vs. GLS Noise vs. Silence

P14
OLWJ vs. Noise MHSN vs. Noise GLS vs. Noise OLWJ vs. MHSN OLWJ vs. GLS

OLWJ vs. Silence MHSN vs. Silence GLS vs. Silence MHSN vs. GLS Noise vs. Silence

Fig. 4. Topographical patterns that exhibit considerable differences in PLI between sound backgrounds within the alpha2 band. The single subjects
P8 and P14 are shown. Solid lines appear only when the absolute values of the corresponding differences are greater than a threshold T. T (= 50%
of the maximum for PLI) was selected to make enough connections visible. For P8, T = 0.3508; for P14, T = 0.3383. Solid lines in red: sound
backgrounds in red higher than backgrounds in blue; solid lines in blue: backgrounds in blue higher than those in red. (For interpretation of the
references to color in this figure legend, the reader is referred to the web version of this article.)

(P = 0.0097), MHSN (P = 0.0131) and GLS
(P = 0.0151). The values of PLI showed increases for
OLWJ (P = 0.0097), MHSN (P = 0.0131) and GLS
(P = 0.0052) in relation to the silence condition.
Fig. 3 shows regional results for the alpha2 frequency,
in which statistical differences in global mean PLI could be
found, to provide further illustration. Instead of multiple
comparisons adjustment, a predetermined fixed
threshold was selected to act as a statistical filter (i.e.,
P < 0.005). Hence the statistical results as shown in
Fig. 3 can merely be regarded as indicative findings. In
relation to the noise and silence conditions, the music
conditions showed an increase in fronto-parietal, fronto-
temporal, centro-parietal, centro-occipital and temporo-
parieto-occipital PLI. Local frontal, central and temporal
synchronizations were also higher in the music
conditions. No obvious distinctions could be found
between the music conditions and between the noise
and silence conditions.
Fig. 4 presents topographical patterns at participant
level, which affords further insight into the network
differences within the alpha2 band. In general, the
differences between the music and noise conditions and
between the music and silence conditions were in the
same direction (i.e., more increases for music conditions
when compared with the noise and silence conditions),
although the individual patterns varied. For the
comparisons between the music conditions and
between the noise and silence conditions, the
differences moved in all possible directions, and
therefore averaged out when combined.
The mean values of the clustering coefficient CW and
the characteristic path length LW are presented in Table
3. There were significant differences in the alpha2
frequency band. The Wilcoxon signed rank test
demonstrated that CW was higher during listening to
OLWJ (P = 0.0023), MHSN (P = 0.0038) and GLS
(P = 0.0013) in comparison with the noise condition. CW
56 J. Wu et al. / Neuroscience 250 (2013) 49–59

showed higher values for OLWJ (P = 0.0032), MHSN
(P = 0.0045) and GLS (P = 0.0038) as compared with
the silence condition. For LW decreases were found
during listening to OLWJ (P = 0.0007), MHSN
(P = 0.0013) and GLS (P = 0.0019) in relation to the
noise condition. LW was shorter for OLWJ (P = 0.0019),
MHSN (P = 0.0038) and GLS (P = 0.0038) than for the
silence condition.
The analysis results of the normalized clustering
coefficient c and the normalized characteristic path
length k are shown in Table 4. Statistical differences
could be observed within the alpha2 band. The pair wise
comparisons revealed lower c in the music conditions
than in the noise condition: for OLWJ (P = 0.0038),
MHSN (P = 0.0027) and GLS (P = 0.0045). c
decreased for OLWJ (P = 0.0045), MHSN (P = 0.0019)
and GLS (P = 0.0038) in comparison with the silence
condition. For k, the values showed lower during
listening to OLWJ (P = 0.0032), MHSN (P = 0.0023)
and GLS (P = 0.0045) in comparison with the noise
condition. k was found to be shorter for OLWJ
(P = 0.0038), MHSN (P = 0.0027) and GLS
(P = 0.0038) in relation to k in the silence condition.
DISCUSSION
In the present study, we utilized PLI and then graph
theoretical measures to investigate the effects of music
on the neural networks of EEG, in different frequency
bands. We observed that functional connectivity, as
expressed by PLI, increased in music conditions
compared with noise and silence conditions in the
alpha2 band. Moreover, network analysis results
demonstrated salient differences between music and
noise conditions and between music and silence
conditions in the alpha2 band: lower normalized

Table 3. Mean clustering coefficients CW and characteristic path lengths LW for five sound backgrounds in six frequency bands

Bands OLWJ MHSN GLS Noise Silence

Mean ± SD Mean ± SD Mean ± SD Mean ± SD Mean ± SD

CW
Delta 0.1471 ± 0.0211 0.1531 ± 0.0177 0.1560 ± 0.0291 0.1708 ± 0.0272 0.1825 ± 0.0236
Theta 0.1111 ± 0.0361 0.1408 ± 0.0511 0.1290 ± 0.0465 0.1073 ± 0.0290 0.0998 ± 0.0260
Alpha1 0.2449 ± 0.0737 0.2562 ± 0.0879 0.2471 ± 0.0786 0.1827 ± 0.0496 0.1712 ± 0.0461
Alpha2 0.2535a,b ± 0.0694 0.2620a,b ± 0.0846 0.2505a,b ± 0.0769 0.1772a ± 0.0491 0.1727b ± 0.0426
Beta 0.1131 ± 0.0357 0.1282 ± 0.0434 0.1234 ± 0.0379 0.0897 ± 0.0254 0.0865 ± 0.0231
Gamma 0.0431 ± 0.0067 0.0430 ± 0.0070 0.0409 ± 0.0057 0.0411 ± 0.0063 0.0402 ± 0.0088

LW
Delta 5.9712 ± 0.7179 5.6502 ± 0.5414 5.7658 ± 0.7500 5.2700 ± 0.7674 4.8995 ± 0.5453
Theta 10.1972 ± 2.0298 8.5594 ± 1.7975 9.5436 ± 2.1770 10.2188 ± 2.1734 10.5633 ± 2.0963
Alpha1 5.4992 ± 1.7259 5.7722 ± 2.0668 5.6632 ± 1.8426 6.5982 ± 1.6138 7.1106 ± 1.8020
Alpha2 5.0650a,b ± 1.5126 5.2506a,b ± 1.6926 5.7379a,b ± 1.5737 6.9551a ± 1.8377 6.8481b ± 1.6147
Beta 10.4645 ± 2.5420 9.6408 ± 2.2002 10.1846 ± 2.8192 12.0906 ± 1.8751 13.2112 ± 3.0676
Gamma 21.7616 ± 2.8352 21.7082 ± 3.0402 22.2068 ± 2.8199 22.4996 ± 2.9063 23.3341 ± 2.8873
For each band, letter superscripts indicate statistically significant differences between backgrounds (Wilcoxon signed rank test, P < 0.005, Bonferroni-corrected).
a
Significant differences between music and noise conditions.
b
Significant differences between music and silence conditions.

Table 4. Mean normalized parameters CW/CW-s and LW/LW-s for five sound backgrounds in six frequency bands

Bands OLWJ MHSN GLS Noise Silence

Mean ± SD Mean ± SD Mean ± SD Mean ± SD Mean ± SD

c = CW/CW-s
Delta 1.1329 ± 0.0255 1.1291 ± 0.0330 1.1135 ± 0.0234 1.1128 ± 0.0252 1.1592 ± 0.0297
Theta 1.1216 ± 0.0262 1.1090 ± 0.0272 1.1350 ± 0.0408 1.1139 ± 0.0282 1.1244 ± 0.0383
Alpha1 1.0961 ± 0.0325 1.0800 ± 0.0295 1.1240 ± 0.0460 1.1304 ± 0.0433 1.1525 ± 0.0479
Alpha2 1.0853a,b ± 0.0303 1.0696a,b ± 0.0292 1.0873a,b ± 0.0264 1.1529a ± 0.0517 1.1652b ± 0.0556
Beta 1.1375 ± 0.0359 1.1557 ± 0.0489 1.1568 ± 0.0472 1.1635 ± 0.0322 1.1685 ± 0.0618
Gamma 1.1909 ± 0.0560 1.1602 ± 0.0434 1.1836 ± 0.0596 1.1869 ± 0.0560 1.1679 ± 0.0602

k = LW/LW-s
Delta 1.0131 ± 0.0169 1.0147 ± 0.0209 1.0227 ± 0.0203 1.0139 ± 0.0200 1.0170 ± 0.0159
Theta 1.0621 ± 0.0257 1.0483 ± 0.0213 1.0727 ± 0.0358 1.0437 ± 0.0212 1.0455 ± 0.0233
Alpha1 1.0483 ± 0.0194 1.0346 ± 0.0204 1.0589 ± 0.0335 1.0618 ± 0.0193 1.0728 ± 0.0241
Alpha2 1.0396a,b ± 0.0190 1.0332a,b ± 0.0181 1.0420a,b ± 0.0195 1.0767a ± 0.0215 1.0833b ± 0.0266
Beta 1.0744 ± 0.0270 1.0760 ± 0.0248 1.0770 ± 0.0329 1.0837 ± 0.0268 1.0829 ± 0.0355
Gamma 1.0684 ± 0.0453 1.0524 ± 0.0405 1.0441 ± 0.0385 1.0329 ± 0.0318 1.0515 ± 0.0273
For each band, letter superscripts indicate statistically significant differences between backgrounds (Wilcoxon signed rank test, P < 0.005, Bonferroni-corrected).
a
Significant differences between music and noise conditions.
b
Significant differences between music and silence conditions.
 J. Wu et al. / Neuroscience 250 (2013) 49–59
clustering coefficient and characteristic path length during
music listening. These variances reflect a tendency of the
functional networks of the brain toward a more random
structure.
During the past decades, music perception has
attracted great attention with respect to its neural
correlates by means of lesion studies and functional
imaging techniques. Most of these studies are primarily
based on the notion of modularity for music processing
(Peretz and Coltheart, 2003), where a module
corresponds to a specific brain area devoted to aspects
of music perception, including musical feature analysis,
auditory memory, auditory scene analysis, and
processing of musical syntax and semantics (Koelsch
and Siebel, 2005; Koelsch, 2011). The neural correlates
of each aspect could be explored with laboratory music,
composed by various manipulations of the
corresponding musical aspect (Peretz and Zatorre,
2005; Ruiz et al., 2009; Schulze et al., 2011). However,
in reality, music is an orderly sequence of musical
attributes, which possess different complicacies. The
present study thus chose the music heard in our daily
lives as auditory stimulus to reconstruct the natural
surroundings in the laboratory, and probed the neural
correlates via functional co-ordination of cortical areas.
The increased degree of synchronization in the alpha
band while listening to music (Table 2) has also been
reported in several EEG studies (Petsche et al., 1997;
Flores-Gutiérrez et al., 2009; Wu et al., 2012). Our
finding provides further support to the notion that music
perception, just like other higher cognitive function,
requires neural processes in multiple cortical areas
working in a co-operative manner (Bhattacharya and
Petsche, 2005). Combining this finding with the results
of power spectral analysis (Table 1), we inferred that
the increased synchronizations under music perception
were not biased by power differences since the
differences in mean PSD among the sound
backgrounds were not significant.
Music perception showed an increase of the
intraregional and interregional synchronizations (Fig. 3),
suggesting that various cortical areas were implicated in
music processing even though these areas were
physically distant. Enhanced synchronous oscillations
among posterior regions may be related to the sensory
stage of the acoustic information processing in
association areas (Flores-Gutiérrez et al., 2009). The
involvement of the left temporal area implied that
musical stimuli activated brain regions which recruited in
language processing. This corroborates the hypothesis
of an intimate relation between music and language,
indicating that overlapping neural resources are involved
in both music and language appraisal (Koelsch and
Siebel, 2005; Flores-Gutiérrez et al., 2007; Koelsch,
2011). Furthermore, functional interactions among task-
relevant areas within the alpha band were strengthened
under music perception. In terms of the framework
proposed by Palva and Palva (2011), this finding offers
evidence for the assumption that alpha rhythmicity plays
a role in active task-relevant processing, instead of the
suppression of task-irrelevant processing.
57
In the network analysis, the most salient results were
an increased CW and a decreased LW in the alpha2 band
while listening to music as compared with the noise and
silence conditions (Table 3). It is noteworthy that the
values of CW and LW depend on the level of PLI besides
network organization. An enhancement of the mean
synchronization strength would give rise to a higher CW
and a shorter LW. A detailed comparison of these
results with a previous study (Wu et al., 2012), in which
CW and LW were compared among music, noise and
silence conditions in the alpha frequency range for the
same threshold, should be undertaken. Wu et al. (2012)
reported that LW was smaller in music perception, with
relatively larger CW. Although the results of both studies
are in agreement, variances in CW and LW cannot be
simply deemed as genuine changes in network
organization since they are possibly ascribed to the
increase of PLI for music listening.
To solve this problem, the normalized measures c and
k were calculated by comparing each original network
with its corresponding random networks. In this way,
both measures were corrected for the changes in mean
PLI between background conditions. We found lower c
and k during music listening in the alpha2 band (Table
4), implying that the brain shifted toward a more random
configuration for music perception. Note that the
normalized clustering coefficient c was quite close to
1.0, which was much smaller than those of several EEG
and MEG studies (Stam et al., 2007a; Rubinov et al.,
2009; Boersma et al., 2011; Douw et al., 2011). It is
probable that, owing to volume conduction, spurious
interdependencies between recorded signals would
result in high evaluations of the clustering coefficients in
previous studies.
Lower clustering c in the music backgrounds could be
explained by an economical principle (Bassett et al.,
2010; Kitzbichler et al., 2011; Bullmore and Sporns,
2012). Densely connected cortical areas or neurons,
characterized by high clustering, incline to be spatial
neighbors and consequently reduce wiring cost.
Therefore decreased clustering indicates higher wiring
cost of functional networks. In addition, path length k
showed lower values in the music conditions, signifying
that the shortest routes between brain regions
decreased so as to improve parallel efficiency of
information transfer (Stam and Reijneveld, 2007;
Bullmore and Sporns, 2012). The decrease in both c
and k could be interpreted as a shift of network
structure toward a more efficient but less economical
configuration. The changes in brain organization in
response to music resemble those in working memory
task performance (Kitzbichler et al., 2011). This
similarity suggests that the music listening task may
demand greater information processing and cognitive
effort.
One of the aims of the present research was to look
for possible differences in network organization between
musical excerpts. However, no differences could be
found between musical stimuli. These might be due to
the fact that the styles of the chosen musical excerpts
were similar.
58 J. Wu et al. / Neuroscience 250 (2013) 49–59
A potential criticism might be that the differences in
synchronization and network measures merely reflected
transient changes between stimuli. We discarded the
first and last 5-s EEG records of each auditory epoch to
avoid transition effects. Furthermore, if the transition
effects had an impact, there should be distinct
differences between stimuli. But in fact it was not true.
The changes in PLI and network configuration
accordingly could not be attributed to transient changes
between stimuli. Also, little difference between the noise
and silence conditions further indicates that the task-
related functional reorganization is brought about by the
musical effect rather than the acoustic effect.
Another possible confusion could be the use of fixed
frequency ranges. There may be individual variability in
any certain frequency. But individually determined filter
setting directly results in a comparison problem, and
moreover, clear boundaries between relevant bands and
their probable functional roles cannot be ascertained
(Bhattacharya and Petsche, 2005). Hence, for simplistic
account, we analyzed the EEG data by employing
standard EEG frequency bands.
It is controversial whether different frequency effects
exist when the alpha band is split into lower and upper
bands. While several studies have observed distinct
reactivity patterns for the two alpha sub-bands, others
have revealed similar patterns (Fink et al., 2005;
Micheloyannis et al., 2009). There is empirical evidence
that functional specialization of different alpha frequency
bands emerges with increasing task demands (Fink
et al., 2005). In this study, the results for both lower and
upper alpha bands were not identical.
CONCLUSION
In summary, our study provides evidence for the
reconfiguration of human brain functional networks while
listening to music. We found that music perception
induced an increased synchronization of cortical regions
and a more random network configuration. The pattern
changes could be detected particularly in the upper
alpha band. Furthermore, differences in network
architecture were not observed between musical stimuli.
These findings imply that music perception leads to an
emergence of a more efficient but less economical
structure and requires more information processing as
well as cognitive effort.
Acknowledgments—The work described in this paper was sup-
ported by National Basic Research Program of China (Grant
No. 2013CB329502) and National Nature Science Foundation
of China (Grant Nos. 61273338 and 31200769).
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