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Expert pianists do not listen: The expertisedependent influence of temporal perturbationon the production of sequential movements
ARTICLE in NEUROSCIENCE · JUNE 2014
Impact Factor: 3.36 · DOI: 10.1016/j.neuroscience.2014.03.058

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Neuroscience 269 (2014) 290–298

EXPERT PIANISTS DO NOT LISTEN: THE EXPERTISE-DEPENDENTINFLUENCE OF TEMPORAL PERTURBATION ON THE PRODUCTION
OF SEQUENTIAL MOVEMENTS
M. C (MARIEKE) VAN DER STEEN, a,b
E. B. D. MOLENDIJK, a E. ALTENMÜLLER a AND
S. FURUYA a*sensorimotor skills. Previous studies provided evidence
for a salient role of auditory feedback in the precise
control of the production of a tone by perturbing auditory
feedback of motor actions such as shifting the pitch of atone (Pfordresher, 2003, 2005; Purcell and Munhall,
2006; Tumer and Brainard, 2007; Munhall et al., 2009;
Maidhof et al., 2010) and delaying the timing of a tone
(Black, 1951; Yates, 1963; Howell and Archer, 1984;
Hashimoto and Sakai, 2003). A key issue in sensorimotor
control research is the neuroplastic adaptation of feedback
control during movements. Several studies haveinvestigated this topic by comparing the eﬀect of pitch
manipulation on vocalization (Jones and Keough, 2008;
Zarate and Zatorre, 2008; Keough and Jones, 2009) andon piano performance (Pfordresher, 2005, 2012;
Katahira et al., 2008) between skilled, unskilled musicians,
and non-musicians. These studies have characterized
distinct diﬀerences in behavioral and neurophysiologicalresponses to the pitch perturbation between theindividuals with diﬀerent skill levels.
The rhythmic feature of acoustic information also
plays a prominent role in ﬁne motor control during
speech and musical performance. Of particularimportance in expressive musical performance is theﬁne-tuned control of the timing of tones (Bhatara et al.,2011; Furuya and Soechting, 2012; Furuya and
Altenmüller, 2013). Evidence for a tight coupling between
the temporal features of acoustic events and motor
actions has been provided by studies that delayed thetiming of the production of a tone in musical performance(Gates et al., 1974; Pfordresher and Palmer, 2002;
Furuya and Soechting, 2010; Pfordresher and Dalla
Bella, 2011). For example, it was shown that delayed
auditory feedback during piano performance augmented
temporal variability of movements (Pfordresher andPalmer, 2002; Pfordresher, 2003; Pfordresher and Dalla
Bella, 2011). This suggests that skilled individuals
use rhythmic information of tones to maintain the precise
timing control of sequential movements. Similarly,
temporal disruptions of movements caused by thedelayed auditory feedback were evident during speech(Howell and Sackin, 2002). In contrast to pitch, however,to our knowledge, no study has examined theexpertise-dependent role of timing information of auditory
feedback in relation to the production of skilled motor
actions.
Neuroimaging studies have demonstrated a
specialized neural network that connects auditory andaInstitute of Music Physiology and Musicians’ Medicine, Universityof Music, Drama, and Media, Emmichplatz 1, 30175 Hanover,Germanyb
Max Planck Research Group ‘‘Music Cognition and Action’’, Max
Planck Institute for Human Cognitive and Brain Sciences,
Leipzig, Germany

Abstract—Auditory information plays an important role inﬁne motor control such as speech and musical performance. The purpose of this study was to assess expertisedependent diﬀerences in the role of temporal informationof auditory feedback in the production of sequential movements. Diﬀerences in motor responses to the transient delay
of tone production during musical performance between
expert pianists and non-musicians were evaluated. Compared to expert pianists, non-musicians showed more pronounced movement disruptions following the delayed
auditory feedback. For example, in response to a perturbation the inter-keystroke interval was prolonged and the
key-press was longer in non-musicians, while the expert pianist marginally shortened both measures. These distinct differences between groups suggest that extensive musicaltraining inﬂuences feedback control in sequential ﬁnger
movements. Furthermore, there was a signiﬁcant positive
correlation between the age at which the expert pianists
commenced their musical training and the amount of disruption. Overall, these ﬁndings suggest that expert pianists
have a higher level of robustness against perturbations
and depend less on auditory feedback during the performance of sequential movements. Ó 2014 IBRO. Publishedby Elsevier Ltd. All rights reserved.

Key words: auditory feedback, sequential movements,
expertise-dependent differences, plasticity, ﬁne motor control.

INTRODUCTION
Speech and musical performance such as singing andplaying an instrument represent highly-sophisticated*Corresponding author. Address: Institute for Music Physiology andMusicians’ Medicine, Hanover University of Music, Drama and Media,
Emmichplatz 1, Hanover 30175, Germany. Tel: +49-(0)511-3100552; fax: +49-(0)511-3100-557.
E-mail address: auditory.motor@gmail.com (S. Furuya).
Abbreviation: IKI, inter-keystroke interval; CD, ﬁnger-key contact
duration; VEL, keystroke velocity.
http://dx.doi.org/10.1016/j.neuroscience.2014.03.058
0306-4522/Ó 2014 IBRO. Published by Elsevier Ltd. All rights reserved.
290

M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

motor regions in musicians’ brains (Bangert and
Altenmüller, 2003; Baumann et al., 2005; Baumann
et al., 2007; Bangert et al., 2006; Lahav et al., 2007;
Zatorre et al., 2007; Engel et al., 2012; Engel et al.,2013). Pianists showed neural activities at the motor
region by merely listening to a short sequence of piano
tones (Bangert et al., 2006). This motor activation
occurred only when listening to a rehearsed piece(D’Ausilio et al., 2006). Overall, these studies suggest thatin trained musicians auditory information evokes bothactivations in auditory areas as well as in brain areas
related to the relevant movements. Perturbations ofauditory feedback during musical performance might
therefore, at least in trained pianists, automatically evoke
motor action. This idea corroborates the neurophysiological ﬁndings of a larger and earlier neural response following the transient pitch perturbation in skilled pianists ascompared to unskilled players (Katahira et al., 2008). In
addition, untrained individuals may focus largely on
movement execution with little listening to tones. Other
research showed that motor action relies largely on sensory feedback information when performing a novel task
(Kawato, 1999; Seidler et al., 2004; Lametti et al.,2012). This raises an alternative possibility in relation to
the eﬀects of perturbed auditory feedback, namely thatthe motor actions of less skilled individuals are more susceptible to auditory perturbation. Consistent with this, thepitch perturbation during vocalizing a single tone yielded
smaller motor disruption in the trained singers than in
the untrained individuals (Jones and Keough, 2008;
Zarate and Zatorre, 2008).
The present study aimed at addressing the role ofexpertise-dependent diﬀerences in the processing of
temporal information of auditory feedback during the
production of sequential movements. Toward this goal,
diﬀerences in motor responses to the transient delay ofthe production of a tone during musical performance
between expert pianists and non-musicians werecompared. Musical performance is an ideal task toapproach this issue with several advantages over
vocalization. First, this task provides technical possibility
for transiently manipulating the timing of a single tone
(Furuya and Soechting, 2010; Pfordresher, 2014). A
transient sensory perturbation allows for purely probing
sensory feedback control, compared to often-applied permanent manipulation of auditory feedback that can cause
adaptation to the perturbation (Houde and Jordan, 1998;
Jones and Keough, 2008). Second, a comparative study291

of musical performance between skilled and unskilledindividuals enables the assessment of the eﬀects of training (Münte et al., 2002; Zarate and Zatorre, 2008; Kleber
et al., 2013), without a possible confounding aging-eﬀect
as a result of development, which may be the case invocalization (MacDonald et al., 2012).

EXPERIMENTAL PROCEDURES
Participants
Ten expert pianists (10 female, 25.2 ± 4.83 yrs, nineright-handed, one bimanual) and 10 non-musicians (sixfemale, 26.6 ± 5.48 yrs, all right-handed) participated in
the study. The expert pianists studied piano playing atmusic conservatories, whereas the non-musicians haveno or very little experience of studying piano playing. All
expert pianists had at least 12 years of extensive pianotraining (range 12–26; mean 20.2; sd 5.16). Six of thenon-musicians had some sort of musical education inplaying an instrument (e.g., recorder/guitar). These six
participants had on average 2.67 years (sd = 1.34) ofexperience. The experimental protocol was approved by
the local ethics committee of the Hannover Medical
University. All participants gave written informed
consent prior to the experiment.
Experimental design
Participants were asked to play a sequence of seventeen
tones requiring the use of all ﬁve digits of the right hand(Fig. 1). The target inter-keystroke interval (IKI) was
400 ms. The participants played on a digital piano (MP9000; Kawai, Krefeld, Germany) that was connected to a
Windows computer. They were provided with themusical score on a computer screen in front of them andwere allowed to practice to familiarize themselves withthe piano and the music selection. The practice session
prior to the experimental trials required the completion ofﬁve correct trials, judged by the experimenter. During
practice, participants played with a metronome (150beats per minute, IKI = 400 ms) in order to playconsistently and accurately at the target tempo withouterroneous keystrokes. Because some of the nonmusicians could not read a score, the experimenter
demonstrated how to play the sequence. Furthermore,
the start position of the thumb was highlighted on the
keyboard, in order to make sure participants used thedesired ﬁngering during the experiment.

Fig. 1. Musical score that was available for the participants during the trials indicating the sequence and appropriate ﬁngering. For explanation
purposes only, the events during which the manipulations could occur are indicated by the dashed ovals. The latter information was not available tothe participants.

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M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

Following the practice session, participants played thetarget melody in either delayed or normal conditions,presented in a randomized order. In delayed conditions,
there was a delay of the timing of the tone production ofeither the 10th or the 12th tone, corresponding with thestroke of the index, or ring ﬁnger, respectively (Fig. 2A).
Tone production was delayed by 120 ms using acustom-made script written in JAVA, running at 1 kHz
for control and recording of MIDI data from the
keyboard. In the normal condition, the participant played
the melody without any auditory perturbation. The
design consisted of three conditions (delay at the strikewith the index or ring ﬁnger, and no delay), times 15
trial repetitions for each condition, being randomly
provided. The randomized order of the 45 trials made it
impossible for the participants to know whether andwhen a delay would occur.
During the experiment, each trial started with thepresentation of four successive tones of a metronome(tempo = 150 beats per minute) as a cue of the target
tempo. The metronome was turned oﬀ automaticallyafter the participant started to play. The participants
were instructed to play as accurately as possible in thetarget tempo. They were also asked to look at themusical score that was displayed on the computer
screen in front of them in order to have them instead oflooking at their hands on the piano keys.

Data analysis
During the experiment, the time each key was depressed
and the time it was released were recorded. In addition,
we also recorded the speed with which each key wasdepressed. Here, MIDI velocities (indicating loudness ofthe tone) provided by the interface ranged from 1 to127. Using these data, the mean and standard deviationof the inter-keystroke interval (from key depression to
key depression), of the keystroke velocity, and of the
ﬁnger-key contact duration (from key depression to key
release) per trial were computed (Fig. 2B, C). Thechanges in keystroke timing and loudness in responseto the delay were deﬁned as the diﬀerence in the
average values of each measure between the delayed
and normal conditions. The trials of the normal condition
functioned as a baseline. Possible eﬀects of exposure to
the delay during the experiment were assessed bycomputing the diﬀerence between the mean of the ﬁrst
and last ﬁve trials for both perturbed ﬁngers. Again thetrials of the normal condition functioned as a baseline.

Statistical analysis
To test for eﬀects of the delayed auditory feedback on thesubsequent movement production, several three-way
mixed-design analyses of variance (ANOVA) (a < 0.05)
with group (expert pianists and non-musicians) as
between subject variable, and perturbed ﬁnger (index
and ring), and event (strikes or intervals before, during
and after the delay, see Fig. 2) as within subject
variables were run. If the assumption of sphericity was
violated, the Greenhouse–Geisser correction wasapplied. T-tests with Bonferroni correction were
performed as post hoc tests. In addition, one-tailed
t-tests (adjusted a = 0.0125) were performed to test
which level signiﬁcantly diﬀered from zero (=nochange). Furthermore, Pearson’s correlations between
the age at which the expert pianists commenced theirmusical training and the diﬀerent outcome measures

A
delay

CD
before delay
VEL
before delayB

IKI 2nd afterdelay

IKI 1st afterdelay

IKI before
delay
CDat delayVEL
at delayCD
1st after delay

IKI 3rd afterdelay
CD
2nd after delay

VEL
1st after delay

VEL
2nd after delay

C

Expert pianists

Non-musicians

100 ms

100 msFig. 2. (A) Schematic representation of the diﬀerent events. Stars reﬂect key-presses. During the orange key-press delayed auditory feedback
could occur. IKI = inter-keystroke interval; CD = ﬁnger-key contact duration; VEL = keystroke velocity. Schematic representation of the groupmean of the inter-keystroke interval (open rectangles) and ﬁnger-key contact duration (ﬁlled rectangles) of the index ﬁnger perturbation for theexpert pianists (B) and the non-musicians (C). The upper staircase reﬂects the normal condition that functioned as a baseline. The lower staircase
shows the delayed condition (orange rectangle). (For interpretation of the references to color in this ﬁgure legend, the reader is referred to the web
version of this article.)293

M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

were calculated to investigate the consequence of earlymusical training on the eﬀect of the delayed auditory
feedback. The analyses were performed with SPSS(IBM SPSS Statistics 21).

values related to the strikes before and after the delay(Fig. 2A). The value related to the keystroke before thedelay functions as a base line of normal movement
production, the other events show the eﬀect of the delayon the movement production.
Results showed signiﬁcant main eﬀects of group(F(1,18) = 16.29, p = 0.001), and event (F(1.30;23.29) =38.731, p < 0.001) for the mean inter-keystroke intervalchange. Pairwise comparisons as post hoc tests (allp < 0.001) revealed that the diﬀerence in group mean
between the delayed and normal conditions was smaller
for expert pianists compared to non-musicians and thatthe ﬁrst inter-keystroke interval after the applied delay
was longer compared to the other intervals. Theinteraction eﬀect between these two independent
variables (F(1.30;23.29) = 22.81, p < 0.001) showed that
this latter eﬀect was most pronounced for non-musicians(Fig. 3A). The one-tailed t-tests revealed that for the
non-musicians the change in mean inter-keystrokeinterval at the ﬁrst inter-keystroke interval after the

RESULTS
Eﬀect of delayed auditory feedback
The mean change in the inter-keystroke interval, theﬁnger-key contact duration, and the velocity at diﬀerent
events related to the delay-manipulation (Fig. 2) were
calculated for both the index and ring ﬁnger
perturbations in order to investigate the eﬀect of delayed
auditory feedback on the accuracy of the subsequent
movement production. The change in standard deviationof all variables was used to assess the eﬀect of thedelayed feedback on the variability, i.e., consistency ofthe same measures. Although the ﬁngering during theevents diﬀered (depending on the index or ring ﬁnger
manipulation) for all measures the events resemble

A Inter-keystroke interval
70

B

****

C

Contact duration

70
expert pianists
non-musicians

Velocity

2.5

*
2

60

60

1.5

difference mean delay - no delay (ms)difference mean delay - no delay (ms)

40

30

20

10

difference mean delay - no delay (MIDI velocity)

50

50

40

30

20

10

0
0

***

-10

***

1

0.5

0

-0.5

-1

-1.5

-2

-10

before delay 1st after 2nd after 3rd after

event

-20
before delay delay 1st after 2nd after

event

-2.5
before delay delay 1st after 2nd after

event

Fig. 3. Signiﬁcant interaction eﬀects of the impact of delayed auditory feedback. Error bars indicate standard error. Blue and red lines represent
expert pianists and non-musicians, respectively. Group times event interaction for the change in (A) inter-keystroke interval, (B) ﬁnger-key contact
duration, and (C) keystroke velocity. ‘⁄’ indicates the value diﬀered signiﬁcantly from zero. Statistical signiﬁcance: ⁄p < 0.01, ⁄⁄p < 0.005,⁄⁄⁄
p = 0.001, ⁄⁄⁄⁄p < 0.001. Corrected a value was 0.0125. See Fig. 2 for the description of the events. (For interpretation of the references to color
in this ﬁgure legend, the reader is referred to the web version of this article.)294

M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

applied delay was signiﬁcantly diﬀerent from zero(t(9) = 6.01, p < 0.001), again indicating a prolongationof the local tempo as a response to the disruption. For theexpert pianists the change at the second interval after thedelay diﬀered signiﬁcantly from zero (t(9) = 5.21,
p = 0.001), indicating a transient acceleration of thetempo. This mean change in inter-keystroke interval( 8.46 ms) was smaller than the threshold of detectability
in timing diﬀerences by experienced listeners (4% ofinterval) (Repp, 1999), suggesting that this change wastoo small to be noticeable (Fig. 3A). The standard deviationof this measure showed a signiﬁcant eﬀect of time(F(2.07;37.27) = 8.903, p = 0.001). There was more variability for the ﬁrst interval after the delay compared to theinterval before and the third interval after the delay(pairwise comparison resp. p < 0.005 and p < 0.001).
For the change in mean ﬁnger-key contact duration
results also showed signiﬁcant main eﬀects of group(F(1,18) = 10.92, p < 0.005), and event (F(1.56;28.10) =10.82, p = 0.001). Again, pairwise comparisons as post
hoc tests revealed that the diﬀerence in contact duration
between the delayed and normal condition was smaller
for pianists compared to non-musicians (p < 0.001).
Contact duration was longer for the keystroke during thedelay compared to the keystroke before (p < 0.05), theﬁrst (p < 0.005), and the second (p < 0.05) keystroke
after the delay. The interaction between these two maineﬀects (F(1.56;28.10) = 6.29, p < 0.01) showed that thiseﬀect was most pronounced for the non-musicians(Fig. 3B). The one-tailed t-tests revealed that for the
non-musicians indeed the mean change in ﬁnger-key
contact duration following the delay was signiﬁcantly
diﬀerent from zero (t(9) = 3.33, p < 0.01), indicating adelayed occurrence of the key-release. For the expert
pianists the change in contact duration during the ﬁrst
keystroke after the applied delay was signiﬁcantly
shorter than zero (t(9) = 4.53, p = 0.001). Again thisshortening was smaller (mean = 13.53 ms) than theperceptual threshold (Fig. 3B).
The interaction eﬀect between group and event on thechange in mean velocity (F(3,54) = 2.81, p < 0.05)
revealed that the expert pianists pressed harder during
the two sequential keystrokes following the delay, while
the non-musicians used less velocity to press the piano
key during the ﬁrst keystroke after the delay (Fig. 3C).
For the standard deviation of this measure across trials,
the main eﬀect of event was found (F(3,54) = 3.08,
p < 0.05). This indicated that the variability of thevelocity of the keystroke related to the delay was larger
than the velocity of the second keystroke after the delay(pairwise comparison as post hoc test: p < 0.05).
Overall, these results indicate that the delayed
auditory feedback mostly inﬂuences the accuracy of themeasures of interest. Participants adjusted to thedelayed feedback relatively quickly, since only
diﬀerences related to the manipulated key-press or theﬁrst interval after the delay were found. Non-musicians
were more disturbed by the delay than expert pianists
as is indicated by the main eﬀect of group and theinteractions. No diﬀerences between the type of
perturbation, i.e. delay on index or ring ﬁnger, were found.

Eﬀect of exposure
To investigate whether participants adjusted their
response to the delayed auditory feedback overrepetitions, a diﬀerence score between the mean of theﬁrst and last ﬁve repetitions of each type ofmanipulation was calculated. Again, the mean andstandard deviation of the inter-keystroke interval, of the
ﬁnger-key contact duration, and of the velocity wereused as outcome measures. Positive values indicatethat the change in mean or standard deviation of theinter-keystroke interval, the ﬁnger-key contact duration,and the velocity was larger during the last ﬁve
repetitions compared to the change during the ﬁrst ﬁve
repetitions of each type of manipulation. Negativevalues suggest that the diﬀerence was more
pronounced in the ﬁrst ﬁve repetitions compared to thelast ﬁve repetitions of the manipulation.
Results showed a main eﬀect of event(F(1.94;34.84) = 3.76, p < 0.05) for the diﬀerence between
the ﬁrst and last ﬁve trials of the change in meaninter-keystroke interval (Fig. 4). This main eﬀect of event
interacted with both group (F(1.94;34.84) = 7.071, p <0.005) and perturbation (F(2.07;37.23) = 6.35, p < 0.005).
Furthermore, there was a signiﬁcant three-way
interaction between the type of perturbation, the event
related to the perturbation and whether the participant
was an expert pianist or non-musician (F(2.07;37.23) =5.44, p < 0.01). Compared to the ﬁrst ﬁve trials, in thelast ﬁve trials non-musicians shortened the length of theinter-keystroke interval of the index ﬁnger on the ﬁrst
interval after the delay, while prolonging the interval of
the index ﬁnger on the second interval after the delay andthe interval of the ring ﬁnger on the third interval after thedelay. The expert pianists seemed to prolong the interkeystroke interval of the ring ﬁnger during the ﬁrst event
after the delay (Fig. 4). The one-tailed t-tests revealed
that only the shortening of the inter-keystroke interval of
the index ﬁnger by the non-musicians was signiﬁcantly
diﬀerent from zero (t(9) = 3.85 p < 0.005) (Fig. 4).
For the diﬀerence between the ﬁrst and last ﬁve trialsof the change in the mean ﬁnger-key contact duration athree-way interaction between the type of perturbation,
the event related to the perturbation and group was
found (F(2.09;37.57) = 3.37, p < 0.05). However, the onetailed t-tests showed that none of the levels yielded tobe signiﬁcantly diﬀerent from zero.
For the standard deviation of the ﬁnger-key contact
duration an interaction eﬀect between perturbation typeand event was evident for the diﬀerence between theﬁrst and last ﬁve trials (F(2.32;41.73) = 3.53, p < 0.05).
Furthermore a three-way interaction between the type of
perturbation, the event related to the perturbation andgroup was found (F(2.32;41.73) = 3.94, p < 0.05) for the
standard deviation of the ﬁnger-key contact duration.
However, all the one-tailed t-tests showed no signiﬁcantdiﬀerences from zero.
Overall, these results indicate that non-musicians
improved their behavior across repetitions of the same
perturbation. The response to the delayed auditory
feedback over repetitions did not diﬀer between thedifference between first and last five trials (ms)M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

295

Mean inter-keystroke interval
40

20

0

-20

-40

**

-60
before delay

1st after

2nd after

3rd after

event
expert index finger delay - no delay
expert ring finger delay - no delay

non-musician index finger delay - no delay
non-musician ring finger delay - no delay

Fig. 4. Signiﬁcant three-way interaction eﬀects between group, event, and manipulation of the eﬀect of exposure for the mean inter-keystrokeinterval. Expert pianists are represented in blue, non-musicians in red. Positive values indicate that the change in mean inter-keystroke interval waslarger during the last ﬁve repetitions compared to the change during the ﬁrst ﬁve repetitions of each type of manipulation. Negative values suggestthat the diﬀerence was more pronounced in the ﬁrst ﬁve repetitions compared to the last ﬁve repetitions of the manipulation. ‘⁄’ indicates the value
diﬀered signiﬁcantly from zero. Statistical signiﬁcance: ⁄p < 0.01, ⁄⁄p < 0.005, ⁄⁄⁄p = 0.001, ⁄⁄⁄⁄p < 0.001. Corrected a value was 0.0125. Error
bars indicate standard error. See Fig. 2 for the description of the events. (For interpretation of the references to color in this ﬁgure legend, the reader
is referred to the web version of this article.)index and ring ﬁnger. For the well-trained expert pianist no
meaningful eﬀects of repetition were found.
Eﬀect of training in expert pianists
As an indicator of the eﬀect of training, it was investigatedwhether the age at which expert pianists started playing
the piano inﬂuenced their response to the delayed
auditory feedback. Therefore, for the expert pianists
Pearson’s correlations between the age of onset and themean and standard deviation of the inter-keystrokeinterval, of the ﬁnger-key contact duration, and of thevelocity for the diﬀerent events were calculated. For the
perturbation on the index ﬁnger three signiﬁcantcorrelations were found. The second interval after delay
was found to be longer for pianists that started their pianotraining later (r = 0.67, p < 0.05). Furthermore, thelength of the ﬁrst and second key press after delay turned
out to be longer if pianists started to play the piano at alater age (resp. r = 0.91, p < 0.001; r = 0.63, p = 0.05).
Overall, these results show larger responses following
the delay in pianists who commenced the training later.
This indicates that the early-started pianists are morerobust against the auditory perturbation compared topianists who started their piano training at a later age.

DISCUSSION
In the present study we assessed diﬀerences in motor
responses to the transient delay of auditory feedback

during musical performance between expert pianists
and non-musicians. Disruptions of temporal features ofﬁnger movements following the transient delay ofauditory feedback for the non-musicians were found.
The movement disruption indicates the integration ofrhythmic information on auditory feedback into
movement production. By contrast, the expert pianists
only showed marginal, not noticeable, rhythmic
disruption of movements following the delayed
feedback, which conﬁrms robust motor control against
external sensory perturbation. One may argue that themusical sequence played in the current experiment, due
to its simple nature, required less eﬀort from the expert
pianists and was perhaps even monotonous to play.
This might postulate that the expert pianists paid lessattention to the task and therefore the auditory
feedback. The lack of any apparent exposure eﬀects on
the keystrokes for the expert pianists contradicts thispossibility, since it indicates that they performed thetask well throughout the whole experiment. The distinct
diﬀerence between these groups suggests the impact ofextensive musical training on feedback control insequential ﬁnger movements. Further evidence in favorof this view was provided by our correlation analysis
across the pianists, which showed smaller movement
disruptions in pianists who commenced musical training
at an earlier age. Similar to our ﬁnding, previous studies
found that perturbed pitch of vocalized sound yielded
smaller motor disruption in trained singers than in
untrained individuals (Jones and Keough, 2008; Zarate

296

M. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

and Zatorre, 2008). It is therefore likely that in both spectral and temporal domains, musical training reduces reliance on auditory feedback control and endows robust
movement production against external perturbation.
Previous studies that investigated hand movements
requiring a novel sensory-motor transformation, forexample, in response to a prism glass or external
perturbing forces, have demonstrated integration of
sensory feedback both into movement execution as wellas into updating the internal representation of thesensory-motor transformation at the early stage oflearning (Kawato, 1999; Shadmehr and Krakauer,2008). With practice, movement production becomes predictive with less integrating sensory feedback (Shadmehr
and Mussa-Ivaldi, 1994). Dysfunctional integration ofauditory feedback into the ﬁnger movements by thenon-musicians in the present study can therefore be dueto the lack of an accurate representation of the dynamics
and acoustics of the piano. In line with this concept, aneurophysiological study has demonstrated the development of a neural network showing stronger links between
auditory and motor cortices through piano practice(Bangert and Altenmüller, 2003). This specialized network
could enable the expert pianists to play in a feed forwardmanner (Bangert and Altenmüller, 2003). In addition, ourﬁnding suggests that despite listening to the perturbed
auditory feedback their special auditory-motor network
did not automatically evoke motor action. This mechanism enables expert pianists to rely on internal sensory
feedback elicited by internal forward model (Ruiz et al.,2009), rather than on external aﬀerent feedback.
The transient delay of auditory feedback yielded anincrease in the inter-keystroke interval for the nonmusicians. This ﬁnding is in agreement with previous
observations of the slowing down of movements byproviding delayed auditory feedback during speech(Black, 1951) and musical performance (Pfordresher,
2003; Pfordresher and Benitez, 2007). The slow-down
supports the beneﬁt that can be elicited by sensory feedback control with longer latencies. By contrast, theexperts did not display any noticeable changes in the local
tempo of movements following the perturbation. Instead,they transiently increased the key-striking velocity. The
lack of such an increase in the non-musicians indicatesthat this motor response is speciﬁc to trained individuals.
It is possible that the stronger keystroke following a perturbation reﬂects the role of somatosensory feedback inthe control of rhythmic movements. A previous study
found a positive correlation between the peak ﬁnger
acceleration at the moment of ﬁnger-key contact andthe temporal accuracy of the subsequent inter-keystrokeinterval during piano performance (Goebl and Palmer,2008). This suggested that a stronger keystroke facilitates
the timing accuracy of the subsequent keystrokes due tofacilitated somatosensory feedback. A stronger keystroke
following the delayed feedback could reﬂect a compensatory strategy of ensuring rhythmic stability of movements.
Indeed, salient roles of somatosensory feedback in ﬁne
motor control have been demonstrated during speech(Nasir and Ostry, 2006, 2008; Ito and Ostry, 2010;
Lametti et al., 2012) and singing (Kleber et al., 2013).

Furthermore, especially for non-musicians the ﬁngerkey contact duration becomes longer in response to thedelayed auditory feedback. As if, the non-musician
waited for the auditory feedback before they let go of
the piano key. This result suggests that the initiation of
the ﬁnger lift of the key is triggered by the auditory
feedback. A study examining the mechanisms
underlying the control of intersegmental dynamics
during reaching movements also showed that feedback
mediated changes decreased the accuracy of
movement reversal when errors in perception occurred(Sainburg et al., 1999).
For the expert pianists both the mean inter-keystrokeinterval as well as the ﬁnger-key contact duration became
shorter in response to the perturbation. However, these
responses were below the perceptual threshold (Repp,
1999), suggesting that they were not noticeable for theexpert pianists. This might reﬂect an implicit compensation for the delay in order to maintain the global tempoof the motor action (Furuya and Soechting, 2010). It isalso possible that as a response to the perturbation, theexpert pianists struck a key slightly earlier in order to highlight the tone (Goebl, 2001), which may well shorten theinter-keystroke interval.
Over trials, the non-musicians displayed a shorter
inter-keystroke interval following the delayed auditory
feedback during strokes with the index ﬁnger. Thisﬁnding indicates short-term learning, since repetition of
the perturbed keystrokes gives rise to less disruption of
movements. A previous study showed development of aneural linkage between the auditory and motor cortices
after 20 min of piano practice (Bangert and Altenmüller,
2003). A formation of an internal representation of auditory-motor transformation might thus aid in decreasing
reliance on auditory feedback for musical performance.
A previous study found that pianists displayed equal
independence of movement control across ﬁngers(Furuya et al., 2011), possibly due to extensive musicaltraining. By contrast, for musically-untrained individuals
a diﬀerence in independent control of movements acrossﬁngers was substantial (Häger-Ross and Schieber, 2000;
Zatsiorsky et al., 2000). These studies reported less independent movement control for the middle and ring ﬁngers
compared to the index and little ﬁngers. Based on theseﬁndings, a larger reliance on sensory feedback during a
keystroke with the ring ﬁnger compared to a keystrokewith the index ﬁnger might be expected for the non-musicians, but not for the expert pianists. Overall, for the
results on the eﬀect of auditory perturbation, there were
no diﬀerences in the disruption between the index andring ﬁngers for both groups (Fig.3). However, theimprovement for the non-musicians over trials appeared
for the index ﬁnger, but not the ring ﬁnger (Fig. 4). Thissuggests superior controllability of the index ﬁnger forthe non-musicians. These ﬁndings propose ﬁnger-dependent reliance on auditory feedback in the ﬁnger movements for non-musicians, but not for expert pianists.
Overall, the ﬁndings of the present study represent
that expert pianists have a higher level of robustness
against perturbations of auditory delayed feedback andthat they depend less on auditory feedback during theM. C (Marieke) van der Steen et al. / Neuroscience 269 (2014) 290–298

performance of sequential movements compared to nonmusicians.
Acknowledgments—The authors thank Marlehn Lübbert for her
help in running the experiment and Naoki Mizumoto for his help
in developing a program that manipulates auditory feedback.
Furthermore, the authors thank Maike Hoﬀ for her valuablecomments on a previous draft of this paper.
This work was supported by funding from the European Community’s Seventh Framework Programme under the EBRAMUS
project – grant agreement n° 238157.

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(Accepted 27 March 2014)(Available online 5 April 2014)