Exp Brain Res (2010) 201:323–330
DOI 10.1007/s00221-009-2044-6
R ES EA R C H A R TI CLE
How and when auditory action eVects impair motor performance
Alessandro D’Ausilio · Riccardo Brunetti ·
Franco Delogu · Cristina Santonico ·
Marta Olivetti Belardinelli
Received: 22 July 2009 / Accepted: 2 October 2009 / Published online: 22 October 2009
© Springer-Verlag 2009
Abstract Music performance is characterized by com-
plex cross-modal interactions, oVering a remarkable win-
dow into training-induced long-term plasticity and
multimodal integration processes. Previous research with
pianists has shown that playing a musical score is aVected
by the concurrent presentation of musical tones. We inves-
tigated the nature of this audio-motor coupling by evaluat-
ing how congruent and incongruent cross-modal auditory
cues aVect motor performance at diVerent time intervals.
We found facilitation if a congruent sound preceded motor
planning with a large Stimulus Onset Asynchrony (SOA
¡300 and ¡200 ms), whereas we observed interference
when an incongruent sound was presented with shorter
SOAs (¡200, ¡100 and 0 ms). Interference and facilita-
tion, instead of developing through time as opposite eVects
of the same mechanism, showed dissociable time-courses
suggesting their derivation from distinct processes. It seems
A. D’Ausilio (&)
DSBTA, Human Physiology Section,
University of Ferrara, via Fossato di Mortara 17/19,
44100 Ferrara, Italy
e-mail: dsllsn@unife.it
A. D’Ausilio · R. Brunetti · F. Delogu · C. Santonico ·
M. O. Belardinelli
ECONA, Interuniversity Centre for Research on Cognitive
Processing in Natural and ArtiWcial Systems,
via dei Marsi 78, 00185 Rome, Italy
F. Delogu · M. O. Belardinelli
Department of Psychology, Sapienza University of Rome,
via dei Marsi 78, 00185 Rome, Italy
R. Brunetti
Università Europea di Roma,
Via Aldobrandeschi 190, 00163 Rome, Italy
that the motor preparation induced by the auditory cue has
diVerent consequences on motor performance according to
the congruency with the future motor state the system is
planning and the degree of asynchrony between the motor
act and the sound presentation. The temporal dissociation
we found contributes to the understanding of how percep-
tion meets action in the context of audio-motor integration.
Keywords Audio-motor integration · Sensory-motor ·
Music · Action planning · Action facilitation ·
Action interference · Action-perception cycle
Introduction
The theory of common coding postulates that when an
action and a percept share a set of features they tend to be
associated (Prinz 1990). According to such hypothesis, the
presentation of a stimulus, previously associated with a par-
ticular action, will automatically prepare the system to pro-
duce the same action (Hommel et al. 2001). This basic
mechanism might be at the basis of behavioural plasticity
connecting arbitrary stimuli to speciWc motor responses.
The link between sensory and motor coordinate systems, on
one hand, helps motor control and the prediction of the con-
sequence of one’s own action (Wolpert and Kawato 1998).
On the other hand it might serve to anticipate the motor
plan necessary to obtain a desired sensory state (Wolpert
and Kawato 1998), to decode other’s action goals (Rizzol-
atti and Craighero 2004), or even to coordinate one’s own
motor execution with other individuals (Sebanz et al.
2006). In this context, musicians are of special interest
because they are extensively trained in a variety of task-
speciWc sensory-motor domains and do often act in coordi-
nation with other musicians.
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Sensory-motor brain plasticity in musicians is the result
of repeated co-occurrences of speciWc actions and the asso-
ciated sensory eVects (Münte et al. 2002). Musicians, in
fact, have proven to be of great interest in the study of how
speciWc training can shape somato-sensory (Elbert et al.
1995), motor (Pascual-Leone et al. 1995; Hund-Georgiadis
and von Cramon 1999), and auditory representations (Pan-
tev et al. 1998), as well as multimodal integration networks
(Stewart et al. 2003). Musicians have also been used as a
model of both long-term structural (Schlaug et al. 1995)
and short-term functional changes in the brain (Rosenkranz
et al. 2007).
A growing number of studies have focused on the mech-
anisms for the integration of auditory and motor informa-
tion (D’Ausilio 2007; Fadiga et al. 2009; Zatorre et al.
2007). Neuroimaging research has recently addressed this
issue from diVerent perspectives. On one hand, it has been
found that motor and premotor activities can be elicited, in
experts, by passive listening to known melodies (e.g.
Haueisen and Knösche 2001; Bangert et al. 2006; Baumann
et al. 2007). On the other hand, several studies have
focused on short-term music training in non-experts. An
EEG study showed increased activity in sensorimotor areas
in naïve participants, both during muted piano playing and
passive listening (Bangert and Altenmüller 2003). Interest-
ingly, a TMS study further demonstrated that passive lis-
tening to a rehearsed piece induces an increased facilitation
at the level of the primary motor cortex, after only 30 min
of practice (D’Ausilio et al. 2006). Recently it was found
that, in non-musicians, premotor activities are elicited by
passive listening of a rehearsed piece, but not by a diVerent
combination of the same notes (Lahav et al. 2007).
In parallel, behavioural studies examined the eVects of
this audio-motor functional link in musicians. A speciWc
experimental procedure has been developed in which the
participant receives a visual instruction to perform an
action (a musical score) while, at the same time, an auditory
stimulus which can be incongruent or congruent with that
action is presented. The impairment or facilitation gener-
ated by the two kinds of stimuli is measured in terms of
reaction time (RT), error rate, and direction of errors (Prinz
1997). Using this procedure (Drost and colleagues 2005a, b,
2007) measured the eVects of congruent and incongruent
piano sounds on musicians’ performance of visually cued
actions on a piano. Drost et al. (2005b) found longer RTs in
the incongruent condition than in the congruent condition
and demonstrated that this eVect occurs at the stage of
motor programming. Drost et al. (2005a) extended the pre-
vious results, obtained with single notes, to chords. Finally,
Drost et al. (2007) showed that this eVect occurs only when
played notes have the timbre of the musical instrument the
single participant is accustomed to play (e.g. guitar notes
triggered by piano strokes do not elicit the eVect).
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Exp Brain Res (2010) 201:323–330
Both neuroimaging and behavioural studies support the
idea that music training induces a speciWc sensory-motor
functional connection. This connection is evidenced by the
emergence of motor activities while listening to rehearsed
musical excerpts (Lahav et al. 2007) and by the eVects on
motor performance induced by listening to plausible action
eVects (Drost et al. 2005a, b). Summing up, converging evi-
dence was found that when a sound-evoked motor repre-
sentation is analogous to that necessary to execute the piano
movement associated to that very sound, the presentation of
sound “A” interferes with the concurrent motor preparation
for producing sound “B”, while facilitating the preparation
of sound “A”. A limitation of the previous studies has been
that they did not consider the audio-motor process as some-
thing developing in time. Sounds were always presented at
the same time as the visual imperative stimulus.
The temporal characteristics of the audio-motor func-
tional connection are of great theoretical importance for
mainly two reasons: (i) cortical and behavioural processing
develops in time, with dissociable functional steps; (ii) the
auditory stimuli, itself, develops in time. Listening to musi-
cal tones (in expert musicians) might trigger a continuous
process of audio-motor transformation, unfolding in time.
This process, when connected to musical execution, can be
conceived as a parallel search in a multidimensional space
of (sensory-motor) features that gradually converge to a
motor solution. In agreement with this idea, neurophysio-
logical studies have shown that the processing of musical
sounds has a precise time course, characterized by speciWc
neural markers (Shahin et al. 2003, 2008; Kuriki et al.
2006; Pizzamiglio et al. 2005). Therefore, one issue in
urgent need for further research is how these behavioural
eVects concerning the interaction between perception and
action develop over time.
Our main experiment tests the prediction that the ampli-
tude (RT diVerences) and direction of eVects (RT reduction
or increase) will depend on the stimulus onset asynchrony
(SOA) between the piano sound and the visually instructed
action on the piano (visuo-motor task). Our hypothesis is
that the motor task will be interfered in diVerent ways,
depending upon the diVerent stages in which the auditory
processing meets the motor programming (manipulated
through diVerent SOAs). We predict three critical time
points (auditory stimulus presented ¡300, ¡200, and
¡100 ms earlier than the visual stimulus), reXecting three
separate processing stages evidenced in recent neurophysi-
ological research (Shahin et al. 2003, 2008; Kuriki et al.
2006; Pizzamiglio et al. 2005). A ¡100 ms SOA should in
principle align the beginning of the visuo-motor task with
component N1 of auditory evoked potentials (AEP). This
component, elicited by single musical tones (as is the case
of our experiments), does not seem to be heavily modulated
by musical expertise but to reXect higher-order spectral
Exp Brain Res (2010) 201:323–330
sound analysis (Shahin et al. 2003). A ¡200 ms SOA
should instead align the beginning of the task with AEP P2
component. This component instead, has been referred as
the Wrst signature of musical expertise, possibly reXecting
the Wrst stage of auditory-motor transformation (Shahin
et al. 2003; Kuriki et al. 2006). Finally, a ¡300 ms SOA
should instead align the beginning of the task with activities
associated to a complete audio-motor transformation (Pizz-
amiglio et al. 2005; Shahin et al. 2008). We hypothesize
that aligning the beginning of the visuomotor task with
these three stages of the auditory-motor continuum will
induce dissociable eVects on RT performance.
Experiment 1
Drost et al. (2005b) suggested that pianists’ playing is inter-
fered with by the concurrent presentation of a piano sound
that conXicts with the visually presented score. The aim of
Experiment 1 was to determine the direction of the eVects
in terms of facilitation or interference by contrasting them
with a NoSound baseline (visuo-motor task).
Method
Fourteen right-handed (assessed with OldWeld’s inventory;
OldWeld 1971) subjects (8 females), aged 22–52 (mean
30.5; SD 9.9), participated in the Wrst experiment. They
were pianists, with at least 5 years of formal training. A
questionnaire assessed the weekly amount of practice
(mean 21.61 h; SD 20.06), years of training (mean
15 years; SD 9.2), and music studies starting age (mean
9.46 years; SD 4.03) for each subject. We assessed that
none of the musicians had absolute pitch through a speciWc
question in the pre-experimental questionnaire. In the train-
ing session we veriWed that all subjects had perfect sight-
reading for simple music scores.
Subjects were presented with both visual and auditory
stimuli. Visual stimuli consisted of 5 scores, each display-
ing two notes on a musical staV (B4 followed by G4, A4,
C5, or D5, notated in G-clef) presented in isolation on a
computer screen (IBM ThinkPad, 15” monitor) placed
50 cm away from the subject. Auditory stimuli consisted of
5 piano sounds, (G4, A4, B4, C5, and D5) with a duration
of 400 ms, presented via headphones (AKG, model K271
Studio) at a comfortable listening level. The piano key-
board was a MIDI controller (M-Audio Keystation 49e)
placed on a table between the computer screen and the sub-
ject. Hand conWguration on the piano keyboard was natural:
thumb was placed on the “G4” key, index Wnger on the
“A4”, middle Wnger on the “B4”, ring Wnger on the “C5”,
and little Wnger on the “D5” key. The procedure did not
require any movement of the hand from this position.
325
Stimulus presentation, conditions, randomization, and
response recording were controlled by a custom-made
script in MAX/MSP, Cycling’74, version 4.6.2 program-
ming environment. The session consisted of 700 trials last-
ing on average 3 s each, plus 5 training trials at the
beginning to accustom the subjects to the task. The experi-
ment was divided into 5 blocks of 140 trials and partici-
pants could rest for several minutes between blocks. Each
trial started with a visual presentation of a score indicating
a “B4” on the staV. The score prompted the subject to play
“B4” on the keyboard with the middle Wnger, which was
followed by the appropriate auditory feedback (piano note
B4). A correct key press also triggered the disappearance of
the score and after a 500 ms blank interval a second score
appeared on the screen, the imperative stimulus. The new
score was shown for 130 ms; it showed the previous “B4”
followed by one of four notes: “G4”, “A4”, “C5”, or “D5”.
At this point the subject had to press the piano key indi-
cated by the second note on the staV as quickly as possible,
by using, respectively, the thumb, index, ring, or little
Wnger. No auditory feedback followed the second key
press.
The design consisted in four visual stimuli £ 5 auditory
stimuli (including NoSound) £ 35 trial repetition, ran-
domly presented. The analysis included three auditory con-
ditions, match, mismatch, and no-sound. In the match
condition the visual imperative stimulus (second note on
the staV in each trial) was accompanied by the correspond-
ing piano sound. In the mismatch condition the sound of
one of the other three possible notes was presented. In the
no-sound condition the subject was presented with the
visual imperative stimulus alone. Subjects’ task was to fol-
low the visual score requests and produce the correct action
on the piano keyboard as quickly as possible. The subjects
were told to ignore the sounds. The dependent measure was
RT, that is the time from the onset of the second score to
the corresponding key press on the MIDI keyboard. Outli-
ers larger than mean §2SD were excluded from further
analysis. Repeated measures analysis of variance
(ANOVA) was performed on the three-level factor “condi-
tion” (match, mismatch, no-sound), and a Duncan’s post
hoc analysis was used to test speciWc comparisons.
Results and discussion
Mean RTs were 464 ms (SE 11.29) for the match condi-
tion, 483 ms (SE 17.57) for mismatch, and 448 ms (SE
8.79) for no-sound. A within-subjects ANOVA revealed a
signiWcant main eVect of “cueing” [F(2,26) = 8.46,
p < 0.001]. Post hoc analysis showed a signiWcant diVer-
ence between match and mismatch conditions (p < 0.05),
with longer RT for mismatch (on average 19.14 ms, SE
8.26) and also a signiWcant diVerence between no-sound
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and mismatch conditions (p < 0.001), with longer RT for
the Mismatch (on average 35.3 ms, SE 10.51). The diVer-
ence between match and no-sound was not signiWcant
(p = 0.27; on average 16.2 ms, SE 6.6). Errors were not
analyzed since subjects performance was almost perfect
in terms of accuracy (>99%).
The results of the Wrst experiment conWrm that a motor
plan can be interfered with a sound that evokes a diVerent
motor representation. However, they also show that a con-
gruent piano sound does not facilitate performance, but it
tends to delay the response, though not by as much as a
mismatching sound does. Therefore, we can see that even a
matching sound does not produce an RT eVect which is sig-
niWcantly diVerent from a baseline with no co-occurring
musical sounds. According to these results, it is inappropri-
ate to speak about facilitation in this case.
Experiment 2
In Experiment 1 the visuo-motor task alone did not diVer
from the cued match condition. This lack of facilitation in
the congruent condition is likely due to the fact that facilita-
tion and interference happen at diVerent moments during
the action planning. In Experiment 2 we directly tested the
hypothesis whether this lack of facilitation is caused by
action planning processes. For this purpose, we manipu-
lated the time interval between the auditory cue and the
visual imperative stimulus while maintaining most of the
relevant features of the previous experiment.
Method
Nine right-handed (assessed with the same method used in
Experiment 1) pianists (4 female; age range 22–42 with
mean 26.8 and SD 6.1) participated in the second experi-
ment. None of them had participated in Experiment 1 nor
were informed of the results of the Wrst study. The same
questionnaire as in Experiment 1 assessed the weekly
amount of practice (mean 14.1 h; SD 11.2), years of train-
ing (mean 11.8 years; SD 3.4), and music studies starting
age (mean 9.1 years; SD 2.4) for each subject. Selection
criteria were the same as those used in Experiment 1.
No musicians had absolute pitch and all subjects demon-
strated to have perfect sight reading for simple music
scores.
Subjects were presented with both visual and auditory
stimuli and asked to play the visually presented notes on a
piano keyboard. The main focus of Experiment 2 was that
of exploring the time deployment of the eVects seen in
Experiment 1. In order to reduce the experimental session
length, we reduced the number of Wnger conditions by
selecting a subset of scores and notes used in Experiment 1.
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Exp Brain Res (2010) 201:323–330
SpeciWcally, since the Wrst experiment showed that index
and thumb RTs are highly comparable, only these two
Wngers were used (thumb and index Wnger RT diVerence,
2 ms; SE 11.12). Thumb, index, and middle Wngers were
always kept in the same natural position. Visual stimuli
consisted of scores “G4”, “A4”, and “B4” presented on the
computer screen. Auditory stimuli consisted of the piano
notes “G4”, “A4”, and “B4”.
Task requirements and procedures for Experiment 2
were the same as in Experiment 1 except for the details
explicitly described. The equipment was the same as in
Experiment 1. Stimuli presentation, conditions, randomiza-
tion, and response recording was controlled by a custom-
made script in MAX/MSP. The experiment consisted of
650 trials lasting on average 3 s each, plus Wve training tri-
als at the beginning. The experimental session was divided
into Wve blocks of 130 trials, to allow the subjects to rest
between blocks. As in Experiment 1, a trial consisted of a
“B4” note presented on the screen that signaled the subject
to press that key on the piano (middle Wnger); the correct
key press was followed by the auditory “B4” and triggered
the disappearance of the visual “B4”. The second score,
presented after a 500 ms blank interval, was visible for
130 ms and showed the initial “B4” followed by one of two
notes: “G4” or “A4”. At this point the subject had to press
the piano key indicated by the second note on the staV as
quickly as possible, by using, respectively, the thumb or
index Wnger. No auditory feedback followed the second key
press.
The design included two variables. The Wrst one, “condi-
tion”, analogously to Experiment 1, included three condi-
tions: match, mismatch, and no-sound. The second variable,
“time”, concerned the timing of the audio stimuli (Fig. 1).
Piano sounds, either congruent or not, could appear in six
time positions relative to the onset of the imperative visual
stimulus: ¡300 (t¡300), ¡200 (t¡200), ¡100 (t¡100), 0 (t0),
+100 (t+100), +240 (t+240) ms. A two-factor repeated mea-
sure ANOVA was run considering “time” and “condition”
as within factors. The dependent measure was RT, as in
Experiment 1. Post hoc comparisons (Duncan’s) were used
to test speciWc comparisons.
Results and discussion
The results are shown in Fig. 1. The analysis revealed sig-
niWcant main eVects of “condition” [F(2,16) = 5.812,
p < 0.05] and “time” [F(5,40) = 7.079, p < 0.0001], as well
as a signiWcant interaction [F(10,80) = 7.137, p < 0.0001].
Post hoc analysis showed a signiWcant diVerence between
mismatch and match for t¡300 (p < 0.0001), t¡200
(p < 0.0001), t¡100 (p = 0.0295) and t0 (p = 0.0159) (aver-
age diVerence in RT between mismatch and match:
t¡300 = 33 ms, t¡200 = 32 ms, t¡100 = 12 ms, t0 = 13 ms,
Exp Brain Res (2010) 201:323–330 327

Match
Paradigm Video Motor Audio NoSound
MisMatch

Motor Response

ms
RT
+240

+100
Imperative
Visual Stimulus Match 0
+ sound (/NoSound) MisMatch
SOA
-100

-200 Fig. 2 Results of Experiment 2. Match (dark grey) and mismatch

(light grey) for each time point (¡300, ¡200, ¡100, 0, +100,
-300 +240 ms). Error bars denote standard error, and asterisks signiWcant
comparisons. The horizontal line represents the RTs for the no-sound
condition, and the light grey thick line shows its standard error
Motor Response
& Associated
Sound

Imperative
Visual
Stimulus
Fixation
Point
Fig. 1 Experiment 2 timeline. Graphical representation of the proce-
dure used in the second experiment. We employed a task very similar
to that of experiment 1. Here we add a varying delay between the
presentation of the second visual imperative stimulus and the audio
presentation, either matching or not with the video
t+100 = 6 ms, t+240 = 2 ms). Mismatch RTs were signiW-
cantly longer than no-sound RTs at t¡200 (p = 0.0291), t¡100
(p = 0.0313), and t0 (p = 0.0025), while match RTs were
shorter than no-sound RTs at t¡300 (p < 0.0001) and t¡200
(p = 0.0003). RTs were faster than in Experiment 1 due to
the use of a 2-choice instead of a 4-choice paradigm. Errors
were not analyzed since subjects’ performance was almost
perfect in terms of accuracy (>99%). Summing up the Wnd-
ings, we can conWrm the results of Experiment 1 at t0.
When imperative stimulus and sound were simultaneously
presented, mismatch caused slower responses than both
match and the no-sound baseline. More interestingly, we
found that the diVerence between match and mismatch was
present only at the early time points (t¡300, t¡200, t¡100, and t0).
SpeciWcally, interference with the motor plan was signiW-
cant at the t¡200, t¡100, and t0 ms time points, while facilita-
tion showed its eVect earlier, for the t¡300 and t¡200 ms
conditions. This pattern of results shows that facilitation
and interference, triggered by action eVects, act within a
small time overlap. Facilitation is visible earlier and dis-
solves around the time when interference is beginning to
inXuence subjects’ RTs Fig. 2.
Experiment 3
In Experiment 2 we showed that varying SOAs induced
dissociable eVects on performance, for congruent and
incongruent conditions. However, it is still possible that, at
least to some extent, the auditory cue can inXuence motor
performance not because of its informative content (con-
gruent or incongruent with the motor plan), but because it
provides an aspeciWc auditory signal able to alert (facilita-
tion) or divert (interference) the subject from the motor
planning depending on the given SOA. In order to exclude
this alternative attentional interpretation of our main result,
in Experiment 3 we tested whether the presentation of a
meaningless sound (not associated to any response) has any
eVect on the visuo-motor task.
Method
Twelve right-handed (assessed with the same method used
in Experiment 1) pianists (3 female; age range 18–43 with
mean 26.8 and SD 7.3) participated in the third experiment.
None of them had participated in Experiment 1 or 2 nor was
informed of the results of these studies. The same question-
naire as in Experiment 1 assessed the weekly amount of
practice (mean 15.6 h; SD 12.3), years of training (mean
14.4 years; SD 9.1), and music studies starting age (mean
12.3 years; SD 4.0) for each subject. Selection criteria were
the same as those used in Experiment 1. No musicians had
absolute pitch and all subjects demonstrated to have perfect
sight reading for simple music scores.
Subjects were presented with both visual and auditory
stimuli and asked to play the visually presented notes on a
piano keyboard. The main focus of Experiment 3 was that
of verifying the inXuence of a task-unrelated sound pre-
sented at diVerent SOA (the same as in Experiment 2) to

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subjects’ performance. Task requirements and procedures
for Experiment 3 were the same as in Experiment 2 except
for few details explicitly described. The equipment was the
same as in Experiment 1 and 2. Stimuli presentation, condi-
tions, randomization, and response recording was con-
trolled by a custom-made script in MAX/MSP.
The experiment consisted of 350 trials lasting on average
3 s each, plus 5 training trials at the beginning. The experi-
mental session was divided into 2 blocks of 175 trials, to
allow the subjects to rest between blocks. Trial timeline,
visual imperative stimuli and subjects’ instructions were
the same as in Experiment 2. The only change was that
instead of a congruent/incongruent piano sound, we pre-
sented a burst of white noise. The white noise stimulus
lasted 400 ms (with a 10 ms linear ramp at the beginning
and 30 ms linear ramp at the end). In a pre-experimental
psychophysical assessment, the white noise level was
adjusted to match the intensity of piano notes used in the
previous experiments. The noise was presented at the same
SOAs used in Experiment 2. Therefore, the timing of the
audio stimuli relative to the onset of the imperative visual
stimulus was: ¡300 (t¡300), ¡200 (t¡200), ¡100 (t¡100),
0 (t0), +100 (t¡100), +240 (t¡240) ms. A no-sound condition
was also included as in Experiment 3 as a reference. The
dependent measure was RT. We ran a two-tailed paired
t test (corrected with the Bonferroni method) between no-
sound and all SOAs to verify whether a noise burst
presented at diVerent time points could aVect performance.
In addition, we ran an independent sample two-tailed t test
between the no-sound conditions of Experiment 2 and 3 in
order to verify whether the two groups were comparable in
terms of general performance.
Results and discussion
The two-tailed paired t test between no-sound and all SOAs
showed no signiWcant diVerence for all SOA (t¡300 vs.
NoSound: t(11) = ¡2.24, p = n.s.; t¡200 vs. NoSound:
t(11) = ¡3.05, p = n.s.; t¡100 vs. NoSound: t(11) = 0.65,
p = n.s.; t0 vs. NoSound: t(11) = 1.81, p = n.s.; t¡100 vs.
NoSound: t(11) = 1.81, p = n.s.; t¡240 vs. NoSound:
t(11) = 0.9, p = n.s.). Mean RT and standard errors for each
condition are shown in Table 1. The no-sound conditions
across Experiment 2 and 3 did not diVer (t(19) = 0.668,
p = n.s.—Mean RT: Experiment 2, 407, SE 6; Experiment 3,
422, SE 19). Errors were not analyzed since subjects’
performance was almost perfect in terms of accuracy
(>99%).
On the whole, our results demonstrate that the mere
presentation of an irrelevant sound (at any SOA from the
imperative stimulus) did not aVect subjects’ performance.
Moreover, the fact that the RTs in the no-sound condition
did not change between Experiment 2 and 3 is an
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Exp Brain Res (2010) 201:323–330
Table 1 Results of Experiment 3
t¡300 t¡200 t¡100 t0 T+100 T+240 NoSound
RT 410 411 424 430 430 425 422
SE 20.1 19.8 18.3 18.6 17.9 18.5 19
RT and standard error values for all conditions in Experiment 3
additional corroboration that the two groups of subjects
have analogous RT performance. Therefore, we can con-
Wrm that the inXuence of sound we found in Experiments
1 and 2 is not related to an aspeciWc attentional eVect, but
it is due to the speciWc musical information that that
sounds transmits.
General discussion
The ideomotor principle posits that actions and eVects
become associated due to their repetitive co-occurrence
(Hommel et al. 2001). The mere presentation of an eVect
activates the associated motor plan, whereas, conversely,
motor preparation aVects the ability to detect and process
the associated sensory stimulus. These simple predictions
have been veriWed in a number of studies, conWrming this
approach as a viable tool in the study of action-perception
integration processes (for a review see Schütz-Bosbach and
Prinz 2007). In this context, experts are a model of over-
learning of these kinds of associations. For example, musi-
cians in the audio-motor domain (Repp and Knoblich 2004;
Keller et al. 2007) and dancers in the visuo-motor domain
(Calvo-Merino et al. 2006) demonstrate a higher motor
awareness within their Weld of expertise. Conversely, it has
been found that musicians (Drost et al. 2005a, b, 2007), as
well as touch typists (Rieger 2004), show motor prepara-
tion when attending to stimuli belonging to the domain of
their motor expertise, such as musical notes or printed let-
ters. However, little is known about the time course of these
latter eVects.
Our experiments speciWcally examined the temporal
deployment of the interactions between sound and move-
ment in audio-motor experts. Moreover, the introduction of
two control conditions, one with the ecological condition of
no-sound and the other with a non musical sound, allowed
us to have a sharper control of the eVects of congruence and
incongruence between cue and imperative stimulus. We
found a strong facilitation when the sound preceded the
imperative visual stimulus by 200–300 ms. DiVerently, the
interference was eVective in a time window running from
200 ms before to the onset of the visual imperative stimu-
lus. Neither interference nor facilitation occurred when the
sound is presented after the onset of the visual imperative
stimulus.
Exp Brain Res (2010) 201:323–330
Musical tones presented at late SOAs (t¡100 and
t¡240 ms) did not elicit any signiWcant eVect on motor
performance. Such result is consistent with a temporal
model of musical tone processing where the Wrst “motor”
stage should fall around 200 ms after stimulus onset
(Shahin et al. 2003; Kuriki et al. 2006). Thus, at positive
SOAs (+100 and +240 ms) the Wrst stage of audio-motor
conversion appears at least 300 ms after the visual impera-
tive stimulus. We interpret this lack of facilitation as if the
sound-evoked motor resonance occurred too late in the
motor planning cascade.
The earliest SOA (t¡300) only showed facilitation and no
interference on RTs. A 300 ms interval has been indicated
to be the time necessary to terminate an audio-motor trans-
formation and preactivate a full motor representation (Pizz-
amiglio et al. 2005; Shahin et al. 2008). A 300 ms interval
is also congruent with the timing of the premotor cortex
activation (Murray et al. 2006; Pizzamiglio et al. 2005),
often reported in investigations of sound-evoked motor
plans (Lahav et al. 2007; Kaplan and Iacoboni 2007), and
of top-down control processes in music behaviours (Shahin
et al. 2008; Koelsch 2006). Thus, the lack of interference
observed at the ¡300 ms SOA might be caused by a top-
down inhibition of the motor plan evoked by the incongru-
ent sound. Thus, we propose that higher-order processes
need at least 300 ms to fully modulate the eVects of the
sound-evoked plan: after this time lapse from the sound
onset, these top-down processes can take advantage of
audio-motor transformation (facilitation) or inhibit it
(therefore avoiding interferences).
The ¡200 ms SOA (t¡200) instead, evidenced both facil-
itation and interference of subjects’ RTs. This result is in
line with our prediction that 200 ms is the time necessary to
complete the Wrst stage of audio-motor conversion (Shahin
et al. 2003; Kuriki et al. 2006). By the time the subject is
visually prompted to play a musical score, the sound analy-
sis has already reached the Wrst critical step in the audio-
motor continuum. However, this stage might reXect an
almost fast process of motor conversion that is not subject
to higher order control and thus elicit both facilitation and
interference.
The ¡100 ms SOA (t¡100) instead evidenced only inter-
ference. This result might indicate that the motor transla-
tion has not been completed, or otherwise we would have
also seen facilitation. This time lag, according to the above-
mentioned neurophysiological experiments, represents the
auditory analysis culminating in a spectral representation of
a complex musical stimulus (Shahin et al. 2003). Therefore,
the interference we observe at this early stage might be of a
diVerent nature, possibly linked with the perceptual cross-
modal interaction between auditory and visual analyses.
Following this rationale, our interference seems to be due
to the cross-modal incongruence between the sound and the
329
imperative stimulus during the bottom-up conversion of
both auditory and visual stimuli into motor plans. This pro-
cess is very fast and it appears to be eVective before the
motor transformation of the sound has been carried out
exhaustively.
Our results show that facilitation and interference have
two diVerent time-courses, but why would we need two
diVerent time deployments for such processes? One possi-
ble explanation might be found by using an ecological
approach. In fact, motor performance in a simple or over-
learned task is usually optimal, and there is no signiWcant
advantage in speeding up its execution. However, interfer-
ence might play an important role in error correction and
online performance tuning, especially for skilled behav-
iours (Maidhof et al. 2009). These considerations become
even more important if we consider that online monitoring
of motor performance is only a part of the picture. For
instance, musicians can also act in strict coordination with
other individuals. In order to do so, the system is necessar-
ily tuned to perform fast online corrections that are driven
by bottom-up mechanisms and rely on shared representa-
tions of actions and perceptions (Schütz-Bosbach and Prinz
2007; Rizzolatti and Craighero 2004; D’Ausilio 2007).
These considerations are of extreme relevance if we con-
sider that in ecological contexts the observer (listener) is
not passively waiting for information—which is the case
for typical experimental paradigms. In natural contexts,
sensory stimulation can also be produced by other’s actions
that are often the result of concerted activities between the
perceiver and one or more individuals (Sebanz et al. 2006).
Conclusion
In this study, we conWrm that potential action eVects are
translated into motor representations, as already demon-
strated in a variety of behavioural, neuroimaging, and
neurophysiological studies (Schütz-Bosbach and Prinz
2007; Rizzolatti and Craighero 2004). Our results also
show that these sensory-motor transformations modulate
behaviour according to their timing and congruence with
the action that will be prepared. Correspondingly, we spec-
ulate that the auditory-motor translation pre-activates the
motor plan that would produce the incoming stimulation,
but also that the eVects on motor performance strictly
depend on the state the motor system will engage in the
near future. The timing analysis allowed us to describe the
processes of action interference and facilitation as pertain-
ing to two temporally dissociable mechanisms that govern
action performance when it has to be adjusted in response
to external events: a Wrst, early, mechanism able to interfere
with the planned action and a second, late, higher level
mechanism able to facilitate the motor planning. It is also
123
330
possible that the sensory-motor resonance could be qualita-
tively diVerent if the observer (listener) has to imitate
action, respond with a complementary action, or simply
understand action (Newman-Norlund et al. 2007).
In this study, we provided a trace of the temporal
dynamics of the mechanisms underlying sensory-motor
integration together with the observation of dissociation
between facilitation and interference. This evidence can
provide new insights on the understanding of the mecha-
nisms of experts’ performance, action monitoring, and
action understanding.
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