Exp Brain Res (2014) 232:3555–3567

DOI 10.1007/s00221-014-4036-4

RESEARCH ARTICLE

Playing beautifully when you have to be fast: spatial and temporal

symmetries of movement patterns in skilled piano performance
at different tempi
Floris T. van Vugt · Shinichi Furuya · Henning Vauth ·
Hans‑Christian Jabusch · Eckart Altenmüller 

Received: 21 October 2013 / Accepted: 5 July 2014 / Published online: 25 July 2014
© Springer-Verlag Berlin Heidelberg 2014

Abstract  Humans are capable of learning a variety of
motor skills such as playing the piano. Performance of
these skills is subject to multiple constraints, such as musi-
cal phrasing or speed requirements, and these constraints
vary from one context to another. In order to understand
how the brain controls highly skilled movements, we
investigated pianists playing musical scales with their left
or right hand at various speeds. Pianists showed system-
atic temporal deviations away from regularity. At slow
tempi, pianists slowed down at the beginning and end of
the movement (which we call phrasal template). At fast
tempi, temporal deviation traces consisted of three peak
delays caused by a thumb-under manoeuvre (which we call
neuromuscular template). Intermediate tempi were a linear
combination trade-off between these two. We introduce and
cross-validate a simple four-parameter model that predicted
the timing deviation of each individual note across tempi
Electronic supplementary material  The online version of this
article (doi:10.1007/s00221-014-4036-4) contains supplementary
material, which is available to authorized users.
F. T. van Vugt (*) · S. Furuya · H. Vauth · H.-C. Jabusch ·
E. Altenmüller  2001; Khalfa et al. 2008; Bhatara et al. 2011). Musicians’
Institute of Music Physiology and Musicians’ Medicine,
University of Music, Drama, and Media, Emmichplatz 1,
30175 Hanover, Germany
e-mail: F.T.vanVugt@gmail.com
F. T. van Vugt 
Lyon Neuroscience Research Center CNRS‑UMR 5292,
INSERM U1028, University Lyon-1, 50 Avenue Tony Garnier,
69007 Lyon, France
H.-C. Jabusch 
Institute of Musicians’ Medicine, Dresden University of Music
Carl Maria von Weber, Leubnitzer Str. 17b, 01069 Dresden,
Germany
(R2  = 0.70). The model can be fitted on the data of indi-
vidual pianists, providing a novel quantification of expert
performance. The present study shows that the motor sys-
tem can generate complex movements through a dynamic
combination of simple movement templates. This provides
insight into how the motor system flexibly adapts to vary-
ing contextual constraints.
Keywords  Generalised motor programmes · Timing ·
Motor skill · Expert musicians · Scale playing ·
Movement effectors
Introduction
Humans are able to learn skills by learning sequences of
movements. Sequential motor behaviours such as speech,
typing a text, or music performance are ways of conveying
information, a crucial aspect of human communication. In
order to perform this function, the skills need to be adapt-
able to changing contexts. For example, playing a musical
sequence at various global and local tempi conveys differ-
ent emotional information to listeners (Dalla Bella et al.
capacity to manipulate tempo thus plays a crucial role in
expressive performance (Goebl and Palmer 2013). Previous
studies focused on variation of the organisation of move-
ment kinematics and muscular activity in relation to tempo
during piano playing (Furuya et al. 2011b, 2012). However,
it is not well understood how temporal features of succes-
sive movement elements (i.e. keystrokes) vary across a
wide range of tempi.
How does the brain implement movements at vari-
ous speeds? Movement production over a wide range of
tempi has been proposed to be controlled by parametric

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modulation of a single temporal movement template or
generalised motor programme (Hollerbach and Flash
1982). For example, when participants learn a tapping
sequence in a certain rhythm, the pattern of timing devia-
tions of the inter-tap intervals is preserved even when
they are asked to reproduce the sequence as fast as possi-
ble (Summers 1975). Such a tempo-invariant motor pro-
gramme allows the brain to simplify the manipulation of
the tempo of movements. A prediction derived from such
model is that temporal features of successive keystrokes
should be invariant across tempi in piano playing. This idea
was supported by an exploratory study that found that the
pattern of inter-onset-intervals (IOIs) of successive key-
strokes were highly similar across different tempi during
playing a certain musical phrase (Repp 1994). Similarly,
in discrete motor tasks such as reaching (Hollerbach and
Flash 1982) and ball throwing with the non-dominant hand
(Hore et al. 2005), spatio-temporal features of the move-
ments were maintained across different speeds.
However, an alternative hypothesis is that variation
of speed involves combining a particular set of distinct
movement templates. This may be computationally costly
because the motor system would have to represent multi-
ple motor templates instead of one (Wolpert and Kawato
1999), but makes the brain flexible enough to cope with
cognitive and biomechanical demands that can vary in rela-
tion to tempo. For example, disruption of movements by
altered auditory feedback in piano playing differed accord-
ing to tempo, implying differential reliance on feedback
control across tempi (Furuya and Soechting 2010). In addi-
tion, when playing some types of musical scales, the thumb
has to rotate while pressing a key in order to move the hand
horizontally (Engel et al. 1997; Furuya et al. 2011a), which
would biomechanically constrain keystrokes in terms of
the hand inertia force that changes with tempo. The tempo-
modulation strategy using a linear sum of some funda-
mental movement patterns predicts variation of movement
characteristics (or more specifically, systematic pattern of
spontaneous temporal deviation) with tempi. Similarly,
expressive timing is thought to change qualitatively with
tempo (Honing 2006, 2007), and the temporal invariance of
successive keystrokes across tempi was violated in piiano
performance (Clarke 1982; Desain and Honing 1994;
Windsor et al. 2001).
The present study approaches this question by investigat-
ing piano scale playing. Piano scale playing has been used
successfully to study expert motor control in musicians
because it is well trained among pianists (Wagner 1971;
MacKenzie and Van Eerd 1990; Jabusch et al. 2004, 2009;
van Vugt et al. 2012, 2013a, b). Additionally, understanding
the scale playing movement is important in itself since it is
frequently used as a metric of severity of musician’s dys-
tonia (Jabusch et al. 2004; van Vugt et al. 2014). We used
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Exp Brain Res (2014) 232:3555–3567
musical scales since they are basic elements of the musical
architecture in classical music as well as in jazz, rock, and
pop music. As a consequence, scale playing represents a fun-
damental aspect of piano technique. Further, the advantage
of studying scale playing over playing other, perhaps more
musical, materials is that the task is clearly defined (play-
ing a scale as regularly as possible), and therefore, it is pos-
sible to directly compare performance between musicians.
Furthermore, scale playing is one of the most well-trained
sequences among pianists’ musical materials. This means
that timing deviations in these materials are not likely due
to spontaneous error. Indeed, pianists were shown to exhibit
systematic timing deviations away from regularity that are
highly individual (van Vugt et al. 2013a, b). Overall, these
timing deviations indicated a slowing down at the begin-
ning and end of the sequence, reminiscent of musical phras-
ing. Furthermore, timing deviation traces showed a series of
delays that could be caused by a thumb-under manoeuvre
during playing the piano scale (Engel et al. 1997; van Vugt
et al. 2012). However, it remains unclear how these patterns
of temporal deviations vary with the speed of playing.
The main question in this paper is how the temporal
deviation pattern changes across the different tempi and
playing directions and hands. Our null hypothesis is that
the temporal deviation pattern is similar across the condi-
tions. An alternative hypothesis is that the temporal devia-
tion pattern changes across the conditions. In particular, we
expect firstly that the pattern of slowing down at the begin-
ning and end of the sequence would dominate at slower
tempi. The reason for this is that when speed demands
are low, the nervous system has more resources available
to create aesthetically pleasing sound or indicate musical
phrasing. Secondly, we expected the delays that could be
caused by a thumb-under manoeuvre to become more pro-
nounced at faster tempi. The reason for this is that at higher
speeds, there is less time for the transitions between key-
stroke movements. Once the motor system is no longer able
to perform the movements in the designated time, delays
will arise whose magnitude will roughly equal the differ-
ence between the available and required time. Thirdly, at
intermediate tempi, we expect to find a trade-off between
these two. Our rationale for this is as follows. If slow-
tempo timing pattern arises from resource availability,
then it will be more pronounced when more time is avail-
able. In other words, the timing pattern should vary para-
metrically with speed. Similarly, if the fast tempo timing
delays are due to the thumb-under manoeuvre, this pattern
become more pronounced with increasing speed demands
(i.e. higher tempo). Since both timing patterns vary para-
metrically with tempo, the intermediate tempi should
show a gradual trade-off between the two. In addition, we
test further hypotheses concerning the temporal control of
scale playing. Do the two hands show comparable temporal
Exp Brain Res (2014) 232:3555–3567 3557
deviation patterns? For example, because of better control
of non-muscular force such as inter-segmental dynamics at
the dominant hand compared to non-dominant hand (Sain-
burg and Kalakanis 2000; Sainburg 2002; Heuer 2007), one
would expect different movement control between the two
hands. Alternatively, it may be computationally advanta-
geous to share motor representations between the hands
when both hands execute the same movement.
Finally, how do temporal deviation traces vary when
movements are executed backwards? That is, could the same
motor pattern be used for a movement that is mirrored in
time? Executing a movement sequence in a different order
might introduce different muscle synergies. This means that
if the brain is able to execute movements backwards in time,
it must have abstracted away from the immediate muscular
instruction. In order to answer this question, we compare the
timing pattern of ascending scales with those of descending
scales, which constitute the same movement mirrored in time.
To test these questions straightforwardly, the present
study developed a linear regression model of timing devia-
tions in piano playing across various tempi, movement
directions (ascending and descending scales) and effectors
(the two hands). This enabled us to evaluate whether vari-
ability across successive keystrokes can be best accounted
for in terms of one or multiple movement templates.
Methods
Participants
Nineteen piano students (ten females, nine males, mean
age 23.7, SD 2.9 years) were recruited from the student
pool at the Hanover Music University. All but one were
right-handed. Seventeen participants took piano as their
main subject, while the other two took organ. Participants
reported no history of neurological disorders and had an
accumulated lifetime keyboard practice of 16.9 (SD 7.5)
1,000 h (except for two pianists who did not specify). The
experiment was approved by the ethical review board of the
University of Music, Drama and Media, Hannover, and the
Medical University Hannover (EG Al_2012_/Go13).
Procedure
Participants played two-octave C-major scales, beginning
with the C (131 Hz) one octave below the middle C and
ending with the C (523 Hz) one octave higher than the mid-
dle C (ascending) and back (descending). For notational
convenience, we will write the notes in this scale as c, d, e,
f, g, a, b, c′, d′, … , c″.
The task of playing a two-octave scale is to press fifteen
adjacent keys serially. Playing the ascending scale with the
right hand, the thumb starts out on the c, and then, the index
and middle finger press the adjacent d and e. Now, the thumb
has to move underneath the index and middle finger to strike
the next key, f. The role of this manoeuvre is to move the posi-
tion of the hand rightward. Now, the index finger moves over
the thumb to strike the g. The middle and ring finger can then
in turn strike the a and b. At this point, another thumb-under
movement is performed to strike the c, at which point, the
entire movement is repeated for the next octave. The left hand
movements are analogue for the descending scale starting with
the thumb on c″. In sum, participants played the scales with
the conventional fingering (123123412312345, where the fin-
gers are numbered from 1 = thumb to 5 = little finger).
Participants played ascending and descending scales inter-
leaved. In between the ascending and descending scales, there
was a short pause of about a second and the pianists made
one additional keystroke on either the note C (65 Hz) or C
(1,047 Hz) with the hand not involved in the scale playing.
This keystroke was used in previous analyses to demarcate
the different trial runs, but was discarded in our current anal-
ysis. The procedure was repeated for 5 different tempi indi-
cated by a metronome at either 40, 80, 120, 160 or 200 BPM.
Furthermore, the procedure was performed for the two hands
in turn. The pianists were instructed to play 4 keystrokes per
metronome beat, which means that the keystroke rate varied
between 2.67 and 13.3 keystrokes/s. Pianists were instructed
to play legato style and to aim for temporal evenness.
Participants played on a MP 9000 MIDI keyboard
(Kawai, Krefeld, Germany).
The keyboard’s digital music interface (MIDI out) sig-
nal was captured on a PC using a commercially available
sequencer software (Musicator Win, version 2.12; Music
Interactive Technology, Bergen, Norway). A metronome
was placed near the pianist so that it was clearly audible
over the sound of the digital piano. No headphones were
used. Loudness was configured as follows. Before the first
participant was recorded, the loudness of the digital piano
was set to perceptually match that of an acoustic piano by
one of the authors who holds a piano performance degree.
The loudness was then kept constant throughout the meas-
urements of all participants. We made the MIDI recordings
used in this study freely available at the following link:
http://dx.doi.org/10.6084/m9.figshare.1108249.
Processing of MIDI data
First, we extracted only the correctly played scales (defined
as those trials on which the pianists pressed the keystrokes
of the C-major scale in the correct order with no omissions
and no additional keystrokes), separating ascending and
descending scales. This yielded 13.5 (SD 1.2) scales for
each pianist, tempo, hand and direction. Across participants,
a total of 1,615 keystroke onsets were discarded because
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they were part of incorrect scales, amounting to 2.1 % of
the recorded material. For the individual pianists, this value
ranged from 0.8 to 10.9 %. The average amount of dropped
onsets was 3.2 % (SD 2.6 %). This amount of discarded
note material did not correlate with the accumulated prac-
tice hours of the pianist (taken as a measure of pianistic
expertise) (Spearman ρ(14) = −.16, p = .57).
Individual note timing
In order to establish the timing of individual notes, we used
the following procedure (van Vugt et al. 2012, 2013b). For
each ascending and descending scale, we computed the
(note, time) pairs, where note is the rank of the note in the
particular scale (0 for c, 1 for d, etc., until 14 for c″) and
time represents the keystroke onset time. Subsequently, we
fitted an ordinary least-square regression line to these pairs.
We rejected fits with an R2 of <0.9 (which occurred when a
scale was played very unevenly), and this was the case for
one scale run (0.02 % of the data). The remaining fits had
an average R2 of 0.9996 (SD 3 × 10−4). The fitted line ena-
bled us to determine the expected onset time for each key-
stroke as the intersection of the regression fit with the hori-
zontal line representing the corresponding note. Finally, we
computed the time difference between this expected onset
and the actual keystroke onset (in msec, with negative dif-
ference reflecting notes played earlier than predicted). We
will refer to this quantity as temporal deviation.
Towards a model of timing deviations
Using the above procedure, we computed the median timing
deviation for each note, hand, playing direction (inward and
outward; inward being defined as radial playing direction,
with the arm moving towards the body; outward as ulnar
playing direction, with the arm moving away from the body)
and tempo across players. We investigated whether the tim-
ing deviation traces for the various tempi were dominated by
a single template or by multiple templates. We hypothesised
to reproduce previous findings of a movement template that
reflects phrasing: a slowing down in the beginning and at the
end of the movement. Additionally, to cope with changing
demands, the traces may be qualitatively different in the con-
text of higher movement production speed. We would expect
such a high-speed template to be dominated by the thumb-
under manoeuvre in outward scales and finger cross-over
manoeuvre in inward scales. We generated two timing tem-
plates that were represented as a vector of timing deviations
for the notes in the scale. We then explored whether the tim-
ing deviations at the intermediate tempi could be rewritten as
a linear combination of these two vectors.
Next, we generalised our model across the hands and
movement directions. First, we cross-validated the model
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Exp Brain Res (2014) 232:3555–3567
fit on the right hand outward scales by assessing its fit on
the left hand outward scales, which are produced by the
same movement but mirrored in space. Second, we turned
to the inward scales for both hands, which are in the same
movement but mirrored in time (that is, played backwards).
That is, we investigated whether the movement templates
that we identified in the steps above can be mirrored in
time (that is, played backwards) or whether they remain
time-invariant.
Fitting methods
For fitting, we used the analytic least-squares solution pro-
vided by ordinary least-squares fitting for generalised lin-
ear models. For nonlinear models that do not allow this, we
instead used the Newton-type optimisation using R’s nlm
function. Furthermore, the nonlinear models with constrained
parameter spaces were implemented using R’s optim func-
tion using the Nelder–Mead simplex method. Whenever
appropriate, we report adjusted goodness-of-fit statistics
2
Radj  = 1 − ((1 − R2) × (N − 1)/(N – k − 1)) where R2 is the
conventional goodness-of-fit, N is the number of observations
and k is the number of parameters in the model. Alternatively,
whenever the variance of the observed data points was impor-
tant, we reported the chi-square measure of goodness-of-fit as
2
follows: χred  = 1/(N – k − 1) × (Σi (Observedi − Predictedi)/
Variancei), where i is the index of the observation, N is the
number of observations and k is the number of parameters in
2 2
the model. The advantage of both these metrics (Radj and χred)
is that they take into account the number of parameters used
in the model and thus prevent overfitting.
In ANOVAs, we report η2G as the generalised effect size
(Bakeman 2005). Whenever Mauchly’s test indicated spheric-
ity violations, we applied the Greenhouse–Geisser correction.
In those cases, for the sake of brevity, we only report the p
value after the correction is applied and marked it as pGG.
Results
Note‑by‑note temporal deviations: establishing the phrasal
and neuromuscular templates
We calculated the note-by-note temporal deviation and
expressed these as a percentage of the interval duration. We
performed a repeated-measures ANOVA with 5 tempi × 2
directions  × 2 hands × 15 notes as factors and temporal
deviation as the dependent variable. There was a main
effect of note (F(14,224)  = 17.49, p < 10−28, η2G  = .16),
reflecting that timing differed across notes. These note-by-
note differences will be the subject of our analysis in the
remainder of this paper. There was no main effect of tempo
(F(4,64) = .49, p = .74) nor of direction (F(1,16) = 3.23,
Exp Brain Res (2014) 232:3555–3567 3559
Fig.  1  a The mean deviation (lateness) in percentage of the inter-
val (right hand outward scales). The depth axis shows the different
tempi. Colour coding is added that reflects the mean vertical position
of each graph segment. The deviation pattern is qualitatively different
p = .09) or hand (F(1,16) = .19, p = .66). Hand and direc-
tion interacted (F(1,16) = 4.93, p = .04) as did tempo and
note (F(56,896) = 8.05, p < 10−48, η2G = .11) and hand and
note (F(14,224)  = 3.30, p < .001, η2G  = .01). This find-
ing constitutes evidence against the generalised motor
programme hypothesis, which would have predicted no
interaction of tempo and note. The other two-way inter-
actions were not significant (both F < .71, p > .58). All
the three-way interactions were significant (all F > 3.72,
p < 10−16) except for tempo, hand and body directions
(F(4,64) = 1.79, p = .14), and finally, the four-way interac-
tion between all factors was significant (F(56,896) = 1.63,
p < .001, η2G  = .01). As a result, we proceeded to analyse
the tempo–note interactions for all four combinations of
hand and direction separately.
First, we focus on the right hand outward scales. We per-
formed a two-way repeated-measures ANOVA with note
and tempo as factors, which yielded a main effect of note
(F(14,224) = 7.95, p < 10−12, η2G = .17), and an interaction
of note and tempo (F(56,896)  = 9.73, p < 10−59), but no
main effect of tempo (F(4,46) = .23, p = .92).
How did note timing vary across tempi? We found that
at fast tempi, three distinct peaks in relative timing appear
(Fig. 1a), corresponding to the positions in the two-octave
scale where the index finger crosses the thumb (e.g., the
index finger playing g after the thumb plays f). At slower
tempi, a different picture emerges: the temporal deviation
trace takes the shape of an arc, with the first few and last
few notes played late, whereas the rest are played slightly
early. Furthermore, at the intermediate tempi, a gradual
trade-off happens between these traces of the extreme
tempi (Fig. 1a). The crucial observation is that variation of
for the slow and fast tempi. b The key down time minus the target
inter-keystroke interval (in msec), revealing that the keys preceding a
thumb passage are released earlier by an amount of time that is more
or less constant across tempi
tempo is achieved as a trade-off between two qualitatively
different timing patterns (templates). This observation
forms the basis of the model of timing deviations that we
provide below. In order to generate this model, in what fol-
lows we (a) established the timing profiles of the fast and
slow tempi, and then (b) parametrised the trade-off between
them across tempi.
Firstly, we generated a simple model of the deviation
trace at the slow tempo (2.7 notes/s). We distinguished two
phases (Fig. 2a): (1) a phase where the temporal deviation
more or less linearly decreases (notes 1:12, i.e. c–g′), i.e.
speeding up in the middle of the scale, and then (2) a phase
where the deviation increases linearly, i.e. slowing down
at the end (notes 13:15, i.e. a′–c″). To model this, we fit
a straight line to the temporal deviation trace from notes c
to g′, and then a second line from a′ to c″. This two-line
2 2
model fitted the data well (Radj   = 0.82, χred(12)  = 0.47)
(Fig. 2c). We took this fit quality based on only two param-
eters as an argument that there was no need to resort to
more complex models of expressive timing to account for
our data. In sum, our model of the slowest tempo implied a
slow start of the scale playing, followed by a smooth speed-
ing up and finishing with a dramatic slowing down. This
finding is consistent with timing arcs found in expressive
playing (Palmer and Krumhansl 1987; Repp 1990; Friberg
and Sundberg 1999; Honing 2003; Friberg et al. 2006) and
is taken to reveal that the two-octave scale is played as a
single phrasal unit (van Vugt et al. 2012). We therefore
refer to this pattern as the phrasal template.
Next, we investigated the deviation trace of the fastest
tempo (13.3 notes/s). Three distinct peaks appeared that cor-
responded to late keystrokes (Fig. 2b). The peaks correspond
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(a)
(c)
Fig.  2  a Mean lateness (expressed as coefficient of variation in %)
for the slowest playing tempo. b Mean lateness for the fastest tempo.
Error bars and shaded area indicate SE across subjects. c Our fit
to the notes played by the index finger, and, to a lesser degree,
the preceding thumb keystrokes. Interestingly, the keystroke
preceding each peak was held down less long, in a way that
did not depend on tempo: the pattern of tone durations was
more or less constant across tempi (Fig. 1b). This suggested
that these timing deviations were dictated by the time the fin-
gers needed to transition between the keystrokes (Engel et al.
1997; Furuya et al. 2011a). This transition time is most prob-
ably due to biomechanical constraints (van Vugt et al. 2013b)
and perhaps also by the hand anatomy (e.g. only the thumb
can perform a three-dimensional rotation that optimises the
horizontal translation of the hand position). Therefore, we
referred to this template as the neuromuscular template. To
construct a model of this template, we proceeded as follows.
We initialised a zero deviation trace where all notes were on
time. Then, we introduced a delay d on the notes played by
the index finger. To partially anticipate for the introduced
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Exp Brain Res (2014) 232:3555–3567
(b)
(d)
to the deviation trace of the slow tempo. The solid line shows our
model. d Our fit at the fastest tempo. The error bars represent the SE
of the mean
delay, the preceding note, played by the thumb, was also
delayed by half that amount, that is, d/2. Finally, the remain-
ing notes were played earlier so that the entire trace centred
around zero. In sum, the index finger was late, the preceding
thumb was half as late and the other notes compensated for
this lateness by being early. This fit is shown in Fig. 2d. This
model had a single parameter, but explained our data well
2 2
(Radj = 0.77, χred(12) = 0.12, p < .001).
Trade‑off between the two templates
Here, we investigate the extent to which we can approximate
the temporal deviation traces of the intermediate tempi as a
combination of the two extreme ones. We computed the lin-
ear combination of the two templates that best fit the devia-
tion trace at each of the tempi. That is, the model contained
10 degrees of freedom (2 parameters per tempo for 5 tempi).
Exp Brain Res (2014) 232:3555–3567 3561
2 2
The overall fit (Radj   = 0.69, χred(62) = 0.43, p < .0001) of
this model was good. However, this fit still contained excess
degrees of freedom, since in this model, an increase in the
contribution of one template did not necessarily entail a
decrease in the other. In order to reduce the parameters to
half, we performed the following fit. Given the deviation trace
dt at a particular tempo as dependent variable, we computed
the fit for dt = α × nm + (1 − α) × phr, where nm and phr
were the (fixed) vectors corresponding to our model fits of the
neuromuscular and phrasal templates, respectively, and α was
(a)
(b)
Fig.  3  a The contributions of the two templates to the timing patterns
across tempi. The squares and their error bars indicate the result of
a least-square fit α(t) × nm + (1 − α(t)) × phr with a single param-
eter (α) for each tempo separately determining a trade-off between
the neuromuscular (nm) and phrasal (phr) templates. Subsequently,
we performed a least-square fit of α(t) = at4 and found the resulting
model, indicated by the solid red line, and 1 −  α(t) is indicated by
the blue line. b Variance explained by the phrasal and neuromuscular
templates for the various tempi on a single-participant basis (see sup-
plementary materials for details) (colour figure online)
the fitted parameter. The corresponding fit had only slightly
2 2
reduced Radj  = .72 (χred(67) = 0.43, p < .0001) relative to the
previous model even though the number of parameters was
divided by two (this latter fit is shown in Fig. 3a). We take this
fact as evidence for a trade-off between the two templates,
rather than free linear combination of the two.
Can one predict how this trade-off parameter (α) varies as a
function of tempo? To this effect, we arbitrarily chose a quartic
function to predict α as a function of tempo as follows. That is,
we fitted α(t) = at4, where t is the tempo (in strokes/s). We
fitted for the parameter a. With this new model, we reduced
the parameters to one (a) instead of four, with a minimal loss
2 2
in explained variance (Radj  = 0.70, χred(70) = 0.40, p < 10−7).
When we fitted a model with only the neuromuscu-
2
lar template, we obtained a much poorer fit (Radj   = 0.40,
2
χred(72) = 0.76, p > .06). The same was true for a model
2
containing only the phrasal template (Radj   = 0.18,
2
χred(71) = 0.55, p < .001). This latter result indicates that
the two templates both contributed to the pattern of timing
deviations observed in these pianists.
In sum, we presented a generative model that predicts
the temporal deviation trace in scale playing based on two
templates that we label as phrasal and neuromuscular and a
trade-off function using a single parameter (a). The phrasal
template is fit using two parameters (the slopes of the two
lines), and the neuromuscular template is based on a sin-
gle parameter (the temporal deviation of the index finger
passage). Using these four parameters, our model predicted
the temporal deviation traces of 5 tempi times 15 notes (75
2
data points). The obtained Radj shows that a considerable
2
portion of the variance is captured (Radj  = 0.70).
Cross‑validation of fit
We tested whether our model of the right hand ascending
scales over-fitted the data. We split the data set in two parts,
the training and test set, in the following way. For each pia-
nist’s hand and playing direction, we allocated the odd trials
to the training set and the even trials to the test set. We then
repeated our analysis on this training set (the odd trials). As
2
with the entire data set, this provided a good fit (Radj  = 0.68,
2
χred(70)  = 0.39, p < .00001; see supplementary materials
for details). Now, we were able to perform the crucial test
of cross-validation: we evaluated this model on the even tri-
als. If our model over-fits the data, we expect it to perform
badly on the even data set. This was not the case: the good-
2 2
ness-of-fit for the even trials (Radj = 0.70, χred (70) = 0.39,
p < .00001) was even better than the fit on the odd trials.
Individual data fit
How robustly can our analysis be applied to individual
pianists? In order to answer this question, we repeated our
13

3562
model fit to the right hand ascending scales for each pianist
individually. Given a pianist and a tempo, we computed the
mean deviation trace (dt) as before, and we approximate
it as a linear combination of the neuromuscular (nm) and
phrasal template (phr) as follows: dt = nm(d) + phr(i, s1,
s2). Here, the neuromuscular model nm was defined by one
parameter (d), and it was constructed precisely as specified
before. Furthermore, the phrasal model was again defined
as two lines, one descending line for notes 1:12 (c–g′) and
one ascending line for notes 13:15 (a′–c″). That is, we
have three parameters: the intercept (i) and slope (s1) of
the first line, and the slope of the second line (s2) (note that
the intercept of the second line is fixed because this line is
forced to intersect with the first one at a′).
That is, we have four parameters for each tempo in the
trace, which we estimate through a nonlinear least-square
fit through the Nelder–Mead simplex method. Addition-
ally, we forced the values of s1 to be negative and d and
s2 to be positive by introducing corresponding penalties in
the squared error term of the optimisation. We then calcu-
lated the amount of variance (in the deviation trace) that is
explained by each of the two templates by subtracting the
summed squared error (SSE) of the two-template model
from the SSE of the other (single) template model. Note
that the difference between this analysis and the trade-off
model (introduced in the previous part of this paper) is that
the latter is based on the average traces (and their SD) of all
participants, whereas the former is fit for each participant
individually.
Figure  3b shows the amount of variance explained by
each template. The result corroborates our previous finding:
the phrasal template explains most variance in the slower
tempi, and the neuromuscular template most of the vari-
ance in the higher tempi. At the intermediate tempi, there is
a clear trade-off (Fig. 3b).
Generalising to all outward scales
In the previous section, we have generated a model for
the timing deviations of the right hand outward scales. As
a cross-validation of our model, we now turn to the left
hand outward scales. When the left hand plays descend-
ing scales, the sequence of movements is exactly the
same as the right hand ascending scales we have ana-
lysed thus far (outward), except in that they are mirrored
in space and executed by a different effector (hand) (van
Vugt et al. 2013b). If the two templates established so far
constitute effector-unspecific representations, then timing
deviations of the left hand outward scales should be simi-
lar to the right hand timing deviations mirrored in space.
Indeed, if we apply the same fit as we have obtained for
the right hand scales to the left hand outward scales, we
obtain a significant model (R2adj  = 0.39, χ2red(70)  = 0.67,
13
Exp Brain Res (2014) 232:3555–3567
p  = .01). Note that the parameters in this model have
been fit to a separate subset of our data (i.e. the right hand
outward scales). Therefore, this evaluation amounts to a
cross-validation (and is actually more stringent than that
because it is evaluated for a different hand). Furthermore,
if we take the outward movements for both hands together
(Fig. 4 right hand panels), our model has a good overall fit
(R2adj = 0.55, χ2red(145) = 0.52, p < 10−7).
Generalising to inward scales
We have presented a model for right hand outward
(ascending) scales and generalised it to left hand outward
(descending) scales. This model shows the brain uses two
motor templates to control both movements. Can the same
motor templates be used to generate the inward scale play-
ing movement (Fig. 4 left hand panels)?
Let us first turn to the fastest tempo. To understand
what temporal deviation trace is predicted for the inward
movement at this tempo, we invite the reader to imagine
the following. If we capture the outward scale on video
and play it backwards, we would see the inward move-
ment. How would the temporal deviation trace for the
original and the backwards variant relate? First of all,
the sequence of notes would be reversed in time. Second,
the temporal deviation of notes would be inverted: notes
that were late in the forward sequence would be early in
the time-mirrored (backwards) sequence and vice versa.
Finally, one further operation is required to complete the
transformation of the timing patterns. We argued previ-
ously that the timing pattern found at the fastest tempo
is determined by the difficulty of the thumb and index
finger transitions. That is, the keystroke after this transi-
tion will be affected (i.e. late) because of the difficulty
of the transition. This means that when the order of the
keystrokes is reversed, the keystroke affected by the
transition delay is the keystroke before the transition. In
other words, we expect the timing pattern of the neuro-
muscular template to be shifted by one note forward in
its entirety. We discard the last note of the inward trace
(since there is no corresponding note in the outward
trace). Since our traces are centred around zero, the tim-
ing of the new data point at the beginning of the trace is
fixed (and it is the same as the discarded note at the end
of the trace).
For clarity, we summarise the process here. We took the
predicted deviation trace of the right hand outward move-
ment at the fastest tempo, reversed the order of the notes,
inverted their temporal deviation (i.e. multiplying the
deviations by −1) and then shifted the entire trace by one
note. The resulting trace correlated highly with the right
hand inward scales [Spearman ρ(14) = .75, p = .001], and,
similarly, for the left hand scales [Spearman ρ(14)  = .73,
Exp Brain Res (2014) 232:3555–3567 3563
p = .001]. Figure 5 illustrates the match between the traces
after this manipulation (compared with Fig. 4 bottom left
and right before this manipulation).
Next, we turn to the slowest tempo, at which we previ-
ously showed the phrasal template dominates. In this case,
inverting the trace in time as shown above does not yield a
good fit for the left hand (ρ(14) = −.34, p = .89) nor for
the right hand (ρ(14)  =  −.63, p  = .99). On the contrary,
the non-inverted, original (time-invariant) traces correlates
highly for the left (ρ(14)  = .75, p < .001) and right hand
(ρ(14)  = .81, p = .0001). In other words, although at the
Fig. 4  The temporal deviation traces for the inward scale movement
(left graphs) and the outward scales movements (right graphs), from
the slowest tempo (top), intermediate tempo (middle) to fastest tempo
(bottom). For simplicity of presentation, we have omitted the other
two measured tempi. The two traces correspond to the left (red) and
fast tempo the timing deviation traces are mirrored in time
(Fig.  4 bottom plots), the timing deviations at the slower
tempo remain invariant (Fig. 4 top plots). Consequently,
the intermediate tempi, which are a linear combination
of these two motor templates, reveal no obvious relation
at face value (Fig. 4 middle plots). Our decomposition of
the timing deviations in two motor templates has revealed
the underlying symmetries in time and space. This finding
shows that hidden symmetries in movements may be read-
ily revealed when movements are decomposed into motor
templates.
right (green) hand. Notes are presented in chronological order, i.e.
note 1 is the first note played, 2 is the note after that. Note how the
traces are similar between the two movements for the slow tempi, but
very different for the faster tempi (colour figure online)
13

3564
Fig. 5  Original outward (red) and re-aligned inward (blue) deviation traces for the fastest tempo. The inward traces are mirrored in time and in
lateness, and shifted by one note (see main text for details). Shaded areas indicate the SE of the mean (colour figure online)
In order to model the inward movements across all
tempi, the predictions were the following. The neuromus-
cular template would invert in time, whereas the phrasal
template remained the invariant. Taking the same param-
eter settings as for the right hand outward scales, we were
able to generate the expected timing pattern, which fit
2
the totality of the inward scale data set well (Radj   = 0.42,
2
χred(149) = .55, p < .001). Again, this amounts to a highly
stringent cross-validation test because the data set on which
the model is evaluated is different (ascending vs. descend-
ing scale playing) than the data set on which the model is
fitted. The fact that the fit is nevertheless significant shows
the robustness of the model.
Generalising to both hands and directions
Finally, we combined the previous models into a single
model for our entire data set of 15 notes across 5 tempi,
2 hands and 2 directions, that is, 300 data points. Using
our model that was fit exclusively on the right hand out-
ward scale, as a cross-validation we generalised to the other
directions and hands as above. The overall model contained
2
four parameters and was highly significant (Radj   = 0.49,
2 −7
χred(299) = 0.53, p < 10 ). Its fit could potentially be fur-
ther optimised by fitting the parameters on the entire data
set, instead of using the parameters fit exclusively on the
right hand outward scales. However, we found the current
analysis more convincing since the remainder of the data
set served as cross-validation.
13
Exp Brain Res (2014) 232:3555–3567
Discussion
The present study aimed to clarify how the brain controls
piano scale playing at various speeds. In particular, how do
temporal deviation patterns in piano scale playing differ
across different tempi and playing directions and hands?
The key prediction of generalised motor programmes (Sha-
piro et al. 1981) is that the timing of motor output scales
with the tempo at which the motor programme is executed.
The present study did not find evidence for this prediction.
Timing deviations were qualitatively different at fast and
slow tempi, as suggested by previous findings (MacKenzie
and Van Eerd 1990). The distributions of the temporal devi-
ations vary strongly between different notes in the scales,
in line with previous findings (van Vugt et al. 2012, 2013b).
We generated a model that was able to explain these varia-
tions as a combination of two qualitatively different timing
templates (i.e., phrasal and neuromuscular templates). Tak-
ing only one of these templates and parametrically modu-
lating it as a function of tempo revealed a significantly infe-
rior model. This observation constitutes evidence against
the relational invariance prediction derived from the gen-
eralised motor programme hypothesis (Repp 1990, 1994).
What do the motor templates identified in this study
represent? During typical playing, the thumb passes
underneath the index and middle finger twice and under-
neath the index, middle and ring finger once (thumb pas-
sages) during the two-octave outward scale playing. After
each such thumb-pass manoeuvre, the index finger has to
Exp Brain Res (2014) 232:3555–3567 3565
pass over the thumb (index passage). At fast tempi, a tem-
plate that instantiates the delays inherent in this so-called
thumb-under manoeuvre (Engel et al. 1997) is dominant.
As a result, we labelled this template as the neuromuscu-
lar template. In conjunction with the finding that tone dura-
tions preceding these delayed keystrokes are systematically
shorter, this template can be explained based on low-level
constraints on finger movements. A fixed amount of time is
required for the fingers to translate to their next keystroke
position. We found that this translation time was independ-
ent of distance: it was the same when the thumb passes
underneath two (between e and f and e′ and f′) or three fin-
gers (between b and c′) (Fig. 2b).
At slower tempi, however, the pattern of deviations
shows two clear landmarks: mildly slower playing in the
beginning and a more pronounced slowing at the end. Tim-
ing traces of this shape are a widespread finding in the
musical timing literature (Palmer and Krumhansl 1987;
van Vugt et al. 2012) known as final ritard (Honing 2003).
They are furthermore reminiscent of timing patterns for
languages across the globe (Turk and Shattuck-Hufnagel
2007). It is similarly a typical feature of locomotion to
slow down at the end in a very similar fashion (Friberg and
Sundberg 1999). That is, we interpret this template as an
expressive device that signals phrasal structure. This inter-
pretation is supported by the finding that this template was
not inverted in time, but remained invariant to temporal
inversion, contrary to the neuromuscular template. The
dominance of the phrasal template at slower tempi reveals
that when temporal constraints on playing are less strict,
pianists shift their cognitive resources to generating an
aesthetically pleasing sound. Furthermore, the fact that the
phrasal template is absent at the fastest tempo suggests that
it reflects not a fundamental feature movement but rather a
control property that is present whenever contextual con-
straints allow it. The phenomenon of slowing down at the
beginning and end appears to be found in action observa-
tion as well. In particular, continuous motion appears to
slow down as a result of our perceptual system’s adaptation
to it (Goldstein 1957). In general, salient events (such as a
start or end of a movement sequence) slow down our per-
ception of time (Tse et al. 2004). However, we argue that
this action observation phenomenon cannot account for the
timing deviations observed in our phrasal template. Pianists
were instructed to aim for temporal evenness, and therefore
if the beginning and end of the scale sounded slower, pia-
nists would have responded with compensatory speeding
up (Penel and Drake 2004).
We observed that two timing templates accounted for
a large portion of the variance in this complex data set of
timing performance across a variety of tempi. We feel this
reflects that the motor system employs a dimensionality
reduction of control (Bernshteı̆n 1967). The motor system
is capable of generating rich repertoires of movements
to fulfil various task demands. This flexibility, however,
causes neural processing to become intractable when hav-
ing to produce fast sequential movements. A simplification
of movement control through combining a small number
of fundamental motor patterns (i.e. primitives) has been
evident in various simple and complex motor behaviours,
such as kicking (Hart and Giszter 2004; d’ Avella and Bizzi
2005), walking (Ivanenko et al. 2004), reaching (d’ Avella
et al. 2006), hand grasping (Santello et al. 2002; Gentner
and Classen 2006) and musical performance (Gentner et al.
2010; Furuya et al. 2011a). Together with the present study,
these findings suggest that the nervous system efficiently
produces a wide repertoire of movements using a small set
of motor primitives.
We found that the timing patterns were similar between
hands and playing directions. This indicated that the brain
was able to generalise motor programmes across the effec-
tors and across the movement directions. Our results indi-
cate a great degree of overlap between motor representa-
tions used to control both the hands (Fig. 4). However,
particularly at faster tempi, the transition-induced hesita-
tions due to the thumb-under manoeuvre appeared to be
more pronounced in the left hand. This is in line with previ-
ous studies revealing inter-manual differences due to inter-
segmental dynamics between the dominant and non-domi-
nant hands (Sainburg and Kalakanis 2000; Sainburg 2002;
Bagesteiro and Sainburg 2002; Heuer 2007).
Production of a planned movement necessitates com-
pensation for neuromuscular constraints on the limb
(Kawato 1999). This requires accurate representation of
the limb dynamics, referred to as an internal model, which
is acquired through extensive training (Thoroughman and
Shadmehr 2000; Osu et al. 2002). However, our finding of
the timing deviations captured by the neuromuscular tem-
plate of skilled pianists indicated failure to compensate for
biomechanical constraints of the hand particularly when
playing at fast tempo. That is, at this speed, pianists did
not play on time on average, but showed systematic timing
irregularities in the form of three peaks of temporal devia-
tion. Why were pianists not able to remove these delays
through extensive practice? The systematic deviations
of the magnitude found here were previously found to be
indistinguishable from perfect regularity even for trained
musicians (van Vugt et al. 2013b). It is thus possible that
the present skilled pianists took maximal advantage of this
perceptual allowance to cope with the biomechanical con-
straints that emerge during the thumb-under movement.
A novel finding is that temporal deviations were main-
tained in movements performed inverted in space (mirrored)
and time (performed backwards). Our study complements
the current understanding of generalised motor programmes
by showing that the motor primitives show a remarkable
13

3566
flexibility in that they can be transferred between effectors
and mirrored in time. Furthermore, the linear combination of
a time-inverted (neuromuscular) and time-invariant (phrasal)
motor template at intermediate tempi results in a motor pattern
that is symmetric neither in space nor in time. This reveals the
sheer complexity of control that can be achieved through sim-
ple combination and mirroring of motor templates.
The model we presented allowed us to adequately
quantify note-by-note timing deviations across a variety
of tempi. The model can be fit to individual pianist’s data
(Fig.  3b). This allows us to gain insight into the relative
prominence of the two templates in a pianists’ playing,
which could constitute a touchstone for metrics of exper-
tise (Jabusch et al. 2009) as well as for a deeper investiga-
tion of anomalies in piano playing in patients with musi-
cian’s dystonia (MD). MD is characterised by structural
and functional maladaptation at cortical and subcortical
regions, which result in a loss of fine motor control (Alten-
müller and Jabusch 2009). Scale playing timing variability
has been shown a meaningful indicator of MD severity in
affected pianists (Jabusch et al. 2004; Rosenkranz et al.
2009; van Vugt et al. 2014) and correlated with neuro-ana-
tomical irregularities (Granert et al. 2011). Based on the
previous findings, we can now ask what qualitative aspects
of playing are altered in MD patients. Furthermore, are the
motor primitives themselves affected or is it their recom-
bination of them that is altered in these patients? The pre-
sent study provides a starting point for tackling these issues
straightforwardly. Furthermore, the current study provides
a basis for identifying symptoms specific to other disorders
such as tendonitis and carpal tunnel syndrome, which have
been prevalent among pianists (Hochberg et al. 1983).
Acknowledgments  This research was supported by the EBRAMUS
(European Brain and Music) Initial Training Network Grant (ITN MC
FP7, GA 238157).
References
Altenmüller E, Jabusch H-C (2009) Focal hand dystonia in musicians:
phenomenology, etiology, and psychological trigger factors. J
Hand Ther 22:144–154. doi:10.1016/j.jht.2008.11.007
Bagesteiro LB, Sainburg RL (2002) Handedness: dominant arm
advantages in control of limb dynamics. J Neurophysiol 88:2408–
2421. doi:10.1152/jn.00901.2001
Bakeman R (2005) Recommended effect size statistics for repeated
measures designs. Behav Res Methods 37:379–384
Bernshteı̆n NA (1967) The co-ordination and regulation of move-
ments. Pergamon Press, Oxford
Bhatara A, Tirovolas AK, Duan LM et al (2011) Perception of emo-
tional expression in musical performance. J Exp Psychol Hum
Percept Perform 921–934. doi:10.1037/a0021922
Clarke EF (1982) Timing in the performance of Erik Satie’s
“Vexations”. Acta Psychol (Amst) 50:1–19. doi:10.1016/
0001-6918(82)90047-6
13
Exp Brain Res (2014) 232:3555–3567
D’ Avella A, Bizzi E (2005) Shared and specific muscle synergies in
natural motor behaviors. Proc Natl Acad Sci 102:3076–3081. doi:
10.1073/pnas.0500199102
D’ Avella A, Portone A, Fernandez L, Lacquaniti F (2006) Con-
trol of fast-reaching movements by muscle synergy combina-
tions. J Neurosci 26:7791–7810. doi:10.1523/JNEUROSCI.
0830-06.2006
Dalla Bella S, Peretz I, Rousseau L, Gosselin N (2001) A develop-
mental study of the affective value of tempo and mode in music.
Cognition 80:B1–B10
Desain P, Honing H (1994) Does expressive timing in music perfor-
mance scale proportionally with tempo? Psychol Res 56:285–
292. doi:10.1007/BF00419658
Engel KC, Flanders M, Soechting JF (1997) Anticipatory and sequen-
tial motor control in piano playing. Exp Brain Res 113:189–199
Friberg A, Sundberg J (1999) Does music performance allude to
locomotion? A model of final ritardandi derived from measure-
ments of stopping runners. J Acoust Soc Am 105:1469–1484.
doi:10.1121/1.426687
Friberg A, Bresin R, Sundberg J (2006) Overview of the KTH rule
system for musical performance. Adv Cogn Psychol 2:145–161
Furuya S, Soechting JF (2010) Role of auditory feedback in the con-
trol of successive keystrokes during piano playing. Exp Brain Res
204:223–237. doi:10.1007/s00221-010-2307-2
Furuya S, Flanders M, Soechting JF (2011a) Hand kinematics of
piano playing. J Neurophysiol 106:2849–2864. doi:10.1152/jn.
00378.2011
Furuya S, Goda T, Katayose H et al (2011b) Distinct inter-joint coor-
dination during fast alternate keystrokes in pianists with superior
skill. Front Hum Neurosci 5:50. doi:10.3389/fnhum.2011.00050
Furuya S, Aoki T, Nakahara H, Kinoshita H (2012) Individual dif-
ferences in the biomechanical effect of loudness and tempo on
upper-limb movements during repetitive piano keystrokes. Hum
Mov Sci 31:26–39. doi:10.1016/j.humov.2011.01.002
Gentner R, Classen J (2006) Modular organization of finger move-
ments by the human central nervous system. Neuron 52:731–742.
doi:10.1016/j.neuron.2006.09.038
Gentner R, Gorges S, Weise D et al (2010) Encoding of motor skill
in the corticomuscular system of musicians. Curr Biol 20:1869–
1874. doi:10.1016/j.cub.2010.09.045
Goebl W, Palmer C (2013) Temporal control and hand movement
efficiency in skilled music performance. PLoS One 8:e50901.
doi:10.1371/journal.pone.0050901
Goldstein AG (1957) Judgments of visual velocity as a function of
length of observation time. J Exp Psychol 54:457–461
Granert O, Peller M, Jabusch H-C et al (2011) Sensorimotor skills
and focal dystonia are linked to putaminal grey-matter volume in
pianists. J Neurol Neurosurg Psychiatry 1225–1231. doi:10.1136
/jnnp.2011.245811
Hart CB, Giszter SF (2004) Modular premotor drives and unit bursts
as primitives for frog motor behaviors. J Neurosci 24:5269–5282.
doi:10.1523/jneurosci.5626-03.2004
Heuer H (2007) Control of the dominant and nondominant hand:
exploitation and taming of nonmuscular forces. Exp Brain Res
178:363–373. doi:10.1007/s00221-006-0747-5
Hochberg FH, Leffert RD, Heller MD, Merriman L (1983) Hand
difficulties among musicians. JAMA J Am Med Assoc
249:1869–1872
Hollerbach MJ, Flash T (1982) Dynamic interactions between limb
segments during planar arm movement. Biol Cybern 44:67–77
Honing H (2003) The final ritard: on music, emotion, and kinematic
models. Comput Music J 27:66–72
Honing H (2006) Evidence for tempo-specific timing in music using a
web-based experimental setup. J Exp Psychol Hum Percept Per-
form 32:780–786. doi:10.1037/0096-1523.32.3.780
Exp Brain Res (2014) 232:3555–3567 3567
Honing H (2007) Is expressive timing relational invariant
under tempo transformation? Psychol Music 35:276–285.
doi:10.1177/0305735607070380
Hore J, O’Brien M, Watts S (2005) Control of joint rotations in
overarm throws of different speeds made by dominant and non-
dominant arms. J Neurophysiol 94:3975–3986. doi:10.1152
/jn.00327.2005
Ivanenko YP, Poppele RE, Lacquaniti F (2004) Five basic muscle acti-
vation patterns account for muscle activity during human locomo-
tion. J Physiol 556:267–282. doi:10.1113/jphysiol.2003.057174
Jabusch H-C, Vauth H, Altenmüller E (2004) Quantification of focal
dystonia in pianists using scale analysis. Mov Disord 19:171–
180. doi:10.1002/mds.10671
Jabusch H-C, Alpers H, Kopiez R et al (2009) The influence of prac-
tice on the development of motor skills in pianists: a longitu-
dinal study in a selected motor task. Hum Mov Sci 28:74–84.
doi:10.1016/j.humov.2008.08.001
Kawato M (1999) Internal models for motor control and trajectory
planning. Curr Opin Neurobiol 9:718–727
Khalfa S, Roy M, Rainville P et al (2008) Role of tempo entrainment in
psychophysiological differentiation of happy and sad music? Int J
Psychophysiol 68:17–26. doi:10.1016/j.ijpsycho.2007.12.001
MacKenzie CL, Van Eerd DL (1990) Rhythmic precision in the per-
formance of piano scales: motor psychophysics and motor pro-
gramming. Atten Perform 13:375–408
Osu R, Franklin DW, Kato H et al (2002) Short- and long-term
changes in joint co-contraction associated with motor learning as
revealed from surface EMG. J Neurophysiol 88:991–1004
Palmer C, Krumhansl CL (1987) Independent temporal and
pitch structures in determination of musical phrases. J Exp
Psychol Hum Percept Perform 13:116–126. doi:10.1037/
0096-1523.13.1.116
Penel A, Drake C (2004) Timing variations in music performance:
musical communication, perceptual compensation, and/or motor
control? Percept Psychophys 66:545–562
Repp BH (1990) Patterns of expressive timing in performances of a
Beethoven minuet by nineteen famous pianists. J Acoust Soc Am
88:622–641
Repp BH (1994) Relational invariance of expressive microstructure
across global tempo changes in music performance: an explora-
tory study. Psychol Res 56:269–284
Rosenkranz K, Butler K, Williamon A, Rothwell JC (2009) Regaining
motor control in musician’s dystonia by restoring sensorimotor
organization. J Neurosci 29:14627–14636. doi:10.1523/JNEURO
SCI.2094-09.2009
Sainburg RL (2002) Evidence for a dynamic-dominance hypoth-
esis of handedness. Exp Brain Res 142:241–258. doi:10.1007/
s00221-001-0913-8
Sainburg RL, Kalakanis D (2000) Differences in control of limb
dynamics during dominant and nondominant arm reaching. J
Neurophysiol 83:2661–2675
Santello M, Flanders M, Soechting JF (2002) Patterns of hand motion
during grasping and the influence of sensory guidance. J Neuro-
sci 22:1426–1435
Shapiro DC, Zernicke RF, Gregor RJ (1981) Evidence for general-
ized motor programs using gait pattern analysis. J Mot Behav
13:33–47
Summers JJ (1975) The role of timing in motor program representa-
tion. J Mot Behav 7:229–241
Thoroughman KA, Shadmehr R (2000) Learning of action through
adaptive combination of motor primitives. Nature 407:742–747.
doi:10.1038/35037588
Tse PU, Intriligator J, Rivest J, Cavanagh P (2004) Attention and
the subjective expansion of time. Atten Percept Psychophys
66:1171–1189
Turk AE, Shattuck-Hufnagel S (2007) Multiple targets of phrase-final
lengthening in American English words. J Phon 35:445–472.
doi:10.1016/j.wocn.2006.12.001
Van Vugt FT, Jabusch H-C, Altenmüller E (2012) Fingers phrase
music differently: trial-to-trial variability in piano scale playing
and auditory perception reveal motor chunking. Front Audit Cogn
Neurosci 3:495. doi:10.3389/fpsyg.2012.00495
Van Vugt FT, Altenmüller E, Jabusch H-C (2013a) The influence of
chronotype on making music: circadian fluctuations in pianists’
fine motor skills. Front Hum Neurosci 7:347. doi:10.3389/fn
hum.2013.00347
Van Vugt FT, Jabusch H-C, Altenmüller E (2013b) Individuality
that is unheard of: systematic temporal deviations in scale play-
ing leave an inaudible pianistic fingerprint. Front Cogn Sci 134.
doi:10.3389/fpsyg.2013.00134
Van Vugt FT, Boullet L, Jabusch H-C, Altenmüller E (2014) Musi-
cian’s dystonia in pianists: long-term evaluation of retrain-
ing and other therapies. Parkinsonism Relat Disord 20:8–12.
doi:10.1016/j.parkreldis.2013.08.009
Wagner C (1971) The influence of the tempo of playing on the rhyth-
mic structure studied at pianist’s playing scales. In: Vredenbregt
J, Wartenweiler J (eds) Med Sport. Karger, Basel, pp 129–132
Windsor L, Aarts R, Desain P et al (2001) The timing of grace
notes in skilled musical performance at different tempi:
a preliminary case study. Psychol Music 29:149–169.
doi:10.1177/0305735601292005
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