Exp Brain Res (2007) 178:363–373
DOI 10.1007/s00221-006-0747-5
R E S E A R C H A RT I C L E
Control of the dominant and nondominant hand:
exploitation and taming of nonmuscular forces
Herbert Heuer
Received: 1 August 2006 / Accepted: 26 September 2006 / Published online: 14 November 2006

Springer-Verlag 2006
Abstract Movements of the dominant and nondomi-
nant hand have been claimed to differ with respect to
how they take intersegmental dynamics into account.
Consistent with this claim, movements of the dominant
hand are hypothesized to better exploit the intrinsic
limb dynamics, whereas movements of the nondomi-
nant hand are controlled to make the intrinsic
dynamics ineffective as far as this is possible. For rapid
finger oscillations this hypothesis implies a higher level
of co-contractions in the nondominant than in the
dominant hand. Replicating previous findings on finger
tapping, finger oscillations of the dominant hand were
faster and less variable than those of the nondominant
hand. More importantly, the variance of the relative
difference between myoelectric signals of antagonistic
muscles and thus the power of reciprocal myoelectric
activity was smaller in the nondominant hand, indi-
cating a relatively higher level of co-contractions than
in the dominant hand. In addition, a spectral decom-
position of the total power of the relative-difference
signal revealed stronger relative power in the fre-
quency band of the finger oscillations in the dominant
than in the nondominant hand. These findings are
consistent with the hypothesis that for the dominant
hand more accurate feedforward control is possible
based on a more accurate internal model of limb
dynamics.
H. Heuer (&)
Institut für Arbeitsphysiologie an der Universität
Dortmund, Ardeystraße 67,
44139 Dortmund, Germany
e-mail: heuer@ifado.de
Introduction
Most humans have a strong preference for using the
right hand in unimanual tasks, a minority prefers the left
hand, and very few people do not exhibit a hand pref-
erence. Whereas hand preference has an asymmetric
U-shaped distribution, hand differences with respect to
skill are normally distributed (Annett 2002, pp 56-66).
Even though the relation between preference and
superiority of performance can be rather complex (cf.
Heuer 1987), it exists in spite of the strikingly different
distributions (Bishop 1989). Therefore the question
arises which factors make the preferred hand more
skilled than the nonpreferred hand, that is, which factors
distinguish the dominant hand (preferred, superior
performance) from the nondominant one (nonpre-
ferred, inferior performance).
This question can be answered in different ways. For
example, manual asymmetries can be related to gross
asymmetries of the brain, in particular to the asym-
metry of language-related areas (Annett 2002). At a
more fine-grained level, the representation of the
dominant hand in the contralateral primary motor
cortex has been shown to be larger than the motor
cortical representation of the nondominant hand
(Hammond 2002; Volkmann et al. 1998). In terms of
functional anatomy, fMRI studies have revealed
stronger activation of the contralateral motor cortex
with paced finger movements of the nondominant than
of the dominant hand (Jäncke et al. 1998). Similarly,
movements of the nondominant hand were associated
with stronger cerebellar activation (Jäncke et al. 1999).
From these and other data it is clear that manual
asymmetries are associated with asymmetries of the
central nervous system, including the spinal cord
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(Melsbach et al. 1996). However, the implications of
these differences for processes of motor control are not
obvious.
With respect to processes of motor control, two
major hypotheses have been posited (Carson 1989).
First, processing of feedback of the dominant hand has
been claimed to be more efficient than processing of
feedback of the nondominant hand. Second, motor
output to the dominant hand has been hypothesized to
be less noisy than output to the nondominant hand.
These hypotheses have been studied primarily for
aimed movements (e.g., Annett et al. 1979; Binsted
et al. 1998; Carson et al. 1993; Roy et al. 1994), and the
evidence turned out to be rather mixed (Carson 1993).
More recently, a new hypothesis has been proposed
by Sainburg (2002), Bagesteiro and Sainburg (2002,
2003) and Sainburg and Kalakanis (2000), called the
dynamic-dominance hypothesis. According to this
hypothesis the dominant arm is more efficient in con-
trolling the effects of intersegmental dynamics, that is,
the effects of those forces that act on a limb segment as
a consequence of the movement of another segment.
For example, Sainburg and Kalakanis (2000) found that
in the dominant arm both movements of the shoulder
joint and of the elbow joint were driven primarily by
muscles acting on the shoulder joint. The rotation of the
elbow joint in this case resulted from the torque due to
the movement of the upper arm. More generally,
interaction torques were more efficiently exploited by
the dominant than by the nondominant arm, and in
addition trajectories of the dominant arm were less af-
fected by interaction torques than those of the non-
dominant arm (Sainburg and Kalakanis 2000). The
superiority of the dominant arm in dealing with inter-
segmental dynamics critically depends on the predict-
ability of the effects of nonmuscular forces. Thus, it is
likely to be based on a more accurate internal model
(cf. Kawato 1999). Compensation of unexpected rather
than predictable inertial loads, in contrast, has been
found to be more efficiently performed with the non-
dominant arm (Bagesteiro and Sainburg 2002).
Although the dynamic-dominance hypothesis is fo-
cussed on intersegmental dynamics, its basic idea can be
generalized to all predictable nonmuscular forces, in
particular to all elements of the intrinsic dynamics of the
moving limb, including inertial forces. With this gener-
alization, the hypothesis holds that movements of the
dominant arm are controlled so that nonmuscular forces
like inertial ones are exploited, which requires that they
can be predicted accurately. In contrast, for movements
of the nondominant arm nonmuscular forces represent a
potential source of disturbances; their effects, therefore,
have to be minimized, that is, the forces have to be
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Exp Brain Res (2007) 178:363–373
tamed as far as this is possible. This hypothesis is con-
sistent with the broader notion of a left-hemisphere
specialization for open-loop control, which depends on
the accuracy of an internal model for the prediction of
nonmuscular forces, and a right-hemisphere specializa-
tion for closed-loop control (cf. Serrien et al. 2006). It is
also consistent with Calvin’s (1983) claim that the tem-
poral precision required by throwing has been critical in
the evolution of the motor mechanisms of the dominant
hemisphere. I shall examine some implications of the
hypothesis for a very simple motor skill, rapid periodic
movements of the index fingers.
Rapid finger tapping is a time-honored task for the
assessment of motor skill (e.g., Ackerman and Cian-
ciolo 2000; Baker et al. 1985; Shimoyama et al. 1990;
Sturm and Büssing 1985; Tinker and Goodenough
1930). Among its main characteristics is a rather con-
sistently found, asymmetry: Tapping is faster and often
less variable with the dominant hand than with the
nondominant hand (Carlier et al. 1993; Elliott et al.
1986; Hammond et al. 1988; McManus et al. 1986;
Peters 1980; Peters and Durding 1979; Provins 1956,
1958; Schmidt et al. 2000; Shimoyama et al. 1990;
Todor and Kyprie 1980). Deviant results of finding no
difference are rare (e.g., Flowers 1975). Peters (1980)
has ascribed the superiority of the dominant hand in
this task to a greater precision of the timing and the
magnitude of the motor outflow.
In rapid finger tapping there is a reactive force,
which results from the impact of the finger on the
surface. The task has also been simplified by way of
omitting the impact, so that rapid oscillations of the
finger (or another limb) have to be produced (e.g.,
Fleishman 1957, 1960; Fleishman and Hempel 1954,
1955). Even in simple oscillatory movements there are
nonmuscular forces, which can be exploited (cf.
Wachholder 1933). In fact, several characteristics of
oscillatory movements are captured by the simplified
mechanical model of a driven damped oscillator (e.g.,
Balera et al. 2000; Hatsopoulos 1996; Heuer 2002,
2006; Heuer and Schulna 2002; Viviani et al. 1976;
Wenderoth and Bock 1999). This simple mechanical
model (or mechanical analogue) offers a useful illus-
tration of the two modes of motor control that exploit
and tame the nonmuscular forces, respectively.
A driven damped oscillator is governed by the dif-
ferential equation
m€x ¼ f
kx
bx_ ¼
kðx
x0 Þ
bx_ ð1Þ
with k and b as stiffness and damping parameters,
respectively. The input f can be written in terms of the
stiffness parameter k and a time-varying target position
Exp Brain Res (2007) 178:363–373 365

x0, so that force is proportional to the deviation of the
actual position from the current target position. Fig-
ure 1 illustrates the results of four simulation runs with
different input signals x0 and different visco-elastic
resistances, that is, different damping and stiffness
parameters. The amplitude of the output signal x is
about 10 units in all four simulations.
The data of Fig. 1a and b illustrate the exploitation
of nonmuscular forces, inertial ones in particular.
Visco-elastic resistance was relatively small (m = 1,
k = 1,000, b = 5). The result is an output signal that is
strongly dependent on the oscillator characteristics. It
is sinusoidal, no matter whether the input signal is
sinusoidal as well (Fig. 1a) or a sequence of pulses
(Fig. 1b), and its amplitude is a multiple of the
amplitude of the input signal. In contrast, the data of
Fig. 1c and d illustrate the taming of nonmuscular
forces. Visco-elastic resistance was relatively large
(m = 1, k = 100,000, b = 500). The result is an output
signal that is tightly coupled to the input. Thus, when
the input signal is sinusoidal, the output signal is
sinusoidal, and when the input signal is a series of
pulses, the output signal is a series of pulses as well.
The amplitude of the output corresponds closely to
that of the input. Note that the input in Fig. 1 is target
or equilibrium position, but not force.
The mechanical characteristics of an oscillating fin-
ger, and visco-elastic resistance in particular, are re-
lated to myoelectric activity. Muscle activation has the
effect of shifting the target position (or equilibrium
position) x0 and of increasing muscle stiffness (Rack
and Westbury, 1969). Independent innervation of
antagonistic muscles provides the means for separate
control of movement and mechanical impedance. Re-
ciprocal activity of flexors and extensors results in
variations of the equilibrium position (or torque). Co-
contraction of antagonistic muscles, in contrast, is
associated with an increase of visco-elastic resistance
(e.g., Hogan 1984; Milner and Cloutier 1998). Thus,
from the present hypothesis, one would expect a higher
level of co-contraction during rapid finger oscillations
of the nondominant hand than of the dominant hand,
so that visco-elastic resistance is increased to tame the
effects of the nonmuscular forces as illustrated in
Fig. 1c and d.
Previous studies of rapid finger oscillations with
the index finger of the dominant hand have revealed
a pattern of alternating flexor and extensor bursts
(Heuer and Schulna 2002; Heuer et al. 2002; Heuer
2006). The phasing of each burst within each move-
ment cycle had immediate kinematic consequences;
an early phasing of a burst resulted in an instanta-

Fig. 1 Simulation results for a) b)

a driven damped oscillator. x x
The upper graph in each part 10 10
of the figure shows the output
x, the lower graph shows the 0 0
input x0, which was sinusoidal
(a, c) and pulse-like (b, d). In -10 -10
a and b visco-elastic
x0 x0
resistance was weak, in c and
d it was strong 10 10

0 0
-10 -10

c) d)
x x
10 10

0 0
-10 -10

x0 x0
10 10

0 0

-10 -10

1s 1s

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366
neous increase of frequency (Heuer and Schulna
2002) and an instantaneous reduction of amplitude
(Heuer 2006). From the present hypothesis, a pattern
of reciprocal activity with the same relation to
kinematic characteristics seems unlikely for the non-
dominant hand, first, because of the expected higher
level of co-contractions, and second, because of the
possibly required smoother modulation of muscular
forces.
Whereas the expected higher level of co-contrac-
tions in the nondominant hand is directly related to the
notion of a style of motor control, which serves to tame
nonmuscular forces, the expectation of a smoother
modulation of muscle forces needs some explanation.
When the movement is dominated by the intrinsic
dynamics of the moving limb, the nature of the mod-
ulation of the muscular forces has few consequences
(cf. Fig. 1a, b); however, the nature of the modulation
of the muscular forces should matter for the nondom-
inant hand with a higher visco-elastic resistance (cf.
Fig. 1c, d). Unfortunately, the consequences of this
hypothesized difference between hands for the myo-
electric signals are ambiguous because of their complex
relation to muscular forces. On the one hand, one
could expect a smoother modulation of myoelectric
activity in the nondominant hand as compared with the
more abrupt or bursting modulation in the dominant
hand, giving rise to a more focussed spectral compo-
sition in the first case. On the other hand, given the
low-pass-filter characteristics of muscles, a smoother
modulation of forces could perhaps also be achieved by
means of higher-frequency small-amplitude modula-
tions of myoelectric activity in the nondominant hand,
which would give rise to a more diffuse spectral com-
position.
From the present hypothesis and the illustrative
example of Fig. 1, one could also expect overall
stronger myoelectric activity in the nondominant than
in the dominant hand. However, this again is a rather
uncertain expectation because of the complex relation
between muscle excitation and tension as well as be-
tween muscle excitation and surface EMG. The rela-
tion between muscular forces and recorded
myoelectric activity cannot be assumed to be the same
for homologous muscles of the dominant and non-
dominant hand. This is true even when myoelectric
activity is normalized (e. g., expressed as percentage of
maximal voluntary contraction). One of the reasons is
that the respective muscles are likely to have different
compositions in terms of different types of muscle fi-
bres. Therefore in the present study total myoelectric
activity is neglected and analyzed only for methodo-
logical reasons given below.
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Exp Brain Res (2007) 178:363–373
Method
Participants
Kinematic and myoelectric data of 20 participants,
aged 18–31 years (mean 22.1 years, SD 4.1 years) were
analyzed. According to the Edinburgh Inventory
(Oldfield 1971), 16 participants were right-handed
(laterality quotients > 70), whereas 4 were left-handed
(laterality quotient < –50). Nine of the 16 right-hand-
ers and 3 of the 4 left-handers were male, the others
were female. Participants volunteered in exchange for
payment. They had given informed consent and could
end their participation in the study whenever they
wanted to. The study was done with the approval of the
institutional ethics committee and in accordance with
the ethical standards laid down in the 1984 Declaration
of Helsinki.
In addition to the data of the 20 participants, which
were actually used for the present study, data were
recorded from additional men and women, but these
were not included in the present analyses for two main
reasons. First, the data of nine additional persons were
not included either because their laterality quotient
was in the range from –50 to +70, so that there was not
a strongly preferred hand, or for reasons of a balanced
design with respect to the order of left-hand trials and
right-hand trials. Second, for 11 additional persons
there had been problems in recording proper myo-
electric signals with modulations at the rate of finger
oscillations, and for 10 persons there were other
problems like perturbations of the recordings or data
loss.
Apparatus
The apparatus was a modification of the one used by
Heuer and Schulna (2002) and Heuer (2006). It was
modified so that it could be used for rapid oscillations
of the left and right index fingers in a horizontal plane.
The hand was held with the thumb upwards and with
the palm resting against a slightly bent surface. It was
supported by a height-adjustable platform. The index
finger was clamped to a light-weight lever of length
6.3 cm and mass 23 g, the pivot of which was aligned
with the metacarpo-phalangeal joint. The lever also
served to immobilize the interphalangeal joints.
The angular position of the index finger was re-
corded by means of an inductive sensor (ID 37/80 with
OD 15-2; TWK-Elektronik, Düsseldorf), which was
connected to the axis of rotation of the lever. Ag/AgCl
electrodes of 2 mm diameter were placed on the
m. flexor digitorum superficialis and the m. extensor
Exp Brain Res (2007) 178:363–373
digitorum of each lower arm according to the guide-
lines given by Zipp (1982). Distance between the
electrodes was 15 mm. The neutral electrodes were
placed above the ulna close to the olecranon of each
arm. Depending on whether the oscillatory movements
were performed with the index finger of the left or
right hand, the electrodes of the left or right arm were
connected to the amplifiers. The EMG signals were
amplified with a gain of 1,000 and band-pass filtered
(10–500 Hz) with a notch filter centered at 50 Hz
(EMG 100 B; Biopac Systems Inc., Santa Barbara). All
signals were recorded at 1,000 samples/s with the
recording system MP 100 WSW (Biopac Systems Inc.).
Design and procedure
Participants were asked to produce maximally rapid
oscillations of the index fingers in a horizontal plane.
Peak-to-peak amplitudes were instructed to be about
30. The 30 range was marked on the apparatus to
allow some visual guidance, but there were no partic-
ular accuracy requirements.
Each test trial lasted 15 s. Participants started and
stopped the rapid finger oscillations upon verbal com-
mands of the experimenter. Breaks between trials
lasted 30 s. Trials were organized in blocks of ten, with
a break of 1 min between blocks. Two blocks of test
trials were performed with the left hand and two with
the right hand. The order of hands was balanced both
for right-handed and left-handed participants. (The
order was almost, but not fully, balanced for male and
female sub-groups.)
Before the first test trial there were a couple of
practice trials (generally 3–5), and again before the
hand was changed. These trials also served to adjust
the electrodes, so that myoelectric activity with a
phasic modulation related to the finger oscillations
could be recorded. Such myoelectric activity could not
always be found, and these cases were not used for the
present study. Therefore the present data are likely to
be a kind of ‘‘positive selection’’ of EMG recordings in
particular of left-hand finger oscillations as far as
phasic EMG activity is concerned.
Data analysis
Figure 2 shows example recordings of 2 s duration.
These are from the last left-hand test trial and the first
right-hand test trial after the change of hands (with two
right-hand practice trials in-between) of a right-handed
male participant. In the first row of graphs angular-
position signals are shown. These were low-pass fil-
tered (fourth-order Butterworth, 20 Hz, dual pass).
367
The second and third row shows the raw EMG signals
of extensor and flexor muscles. Whereas for the right
hand there were well-defined alternating extensor and
flexor bursts, for the left hand the bursts were not that
well-defined, though there was a clearly recognizable
modulation of myoelectric activity. The myoelectric
signals were rectified and smoothed by means of a
moving average of window width 21 ms. The resultant
signals are shown in the fourth and fifth row of Fig. 2.
Note that the scaling of the myoelectric signals is
specific to each graph. The amplitudes of these signals
cannot be meaningfully compared across hands (nor
across muscles) because of their dependence on details
of electrode placement, thickness of tissue above the
muscles etc.
The recordings of each test trial were truncated to
eliminate eventual start-up effects and periods of final
slowing; of the 15 s of each trial the first 2 s and the last
1 s were neglected. For the angular-position signal
peak extensions and peak flexions were identified.
From the times and locations of these landmarks the
durations of cycles defined by successive peak exten-
sions as well as by successive peak flexions were
determined, in addition the amplitudes of extensions
and flexions. Across the cycles of each trial medians
were computed; as a measure of variability the semi-
inter-quartile range was used. Whereas the medians of
the durations of peak-extension and peak-flexion cy-
cles are essentially identical, the measures of variability
can differ. The same is true for the medians and semi-
inter-quartile ranges of flexion and extension ampli-
tudes. For each trial the flexion- and extension-related
measures of cycle duration and amplitude (median and
semi-inter-quartile range in each case) were averaged.
Even though the total myoelectric activity of left-
hand and right-hand muscles cannot meaningfully be
compared for the purpose of the present study, we
determined total myoelectric activity for flexors and
extensors by the integrals of the rectified and smoothed
EMG signals across the 12 s of each trial. The reason
for doing so was to be able to roughly assess eventual
differences between myoelectric signals of both hands
with respect to the level of noise, which is likely to be
higher for weaker signals (cf. Gerdle et al. 1999).
The assessment of co-contractions versus reciprocal
contractions of the antagonistic muscles was based on a
relative-difference signal. The rectified and smoothed
EMG signals were scaled to a mean of 1, and for each
point in time the relative difference (E – F)/(E + F)
was computed, where E and F denote the values of the
rectified, smoothed, and scaled myoelectric signals re-
corded from the extensor and the flexor, respectively.
Examples of the relative-difference signal are shown in
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368 Exp Brain Res (2007) 178:363–373

Fig. 2 Examples of raw and left hand right hand

processed signals for the left
and right hand. The topmost
40°
graphs show angular positions
of the index fingers. The next
two rows of graphs show raw
EMG signals of flexors and
extensors, respectively, and 0.08 0.4
the fourth and fifth row of 0.04 0.0
0.00
graphs shows the rectified and -0.04 -0.4
smoothed EMG signals (on -0.08 -0.8
the ordinates voltages are
shown in mV). The bottom 0.2 0.4
row of graph shows the 0.1 0.2
0.0 0.0
relative-difference signals -0.1 -0.2
-0.2 -0.4

0.02 0.2

0.01 0.1

0.00 0.0

0.10
0.04
0.02 0.05

0.00 0.00

1.0 1.0

0.0 0.0

-1.0 -1.0
0.5 s

the bottom row of graphs of Fig. 2. Values around zero
indicate dominant co-contraction, whereas values close
to +1 and –1 indicate a dominant reciprocal activity.
The pattern of alternating flexor and extensor bursts of
the right hand results in a relative-difference signal
which tends to jump between values close to ±1,
whereas the pattern of myoelectric activity seen in the
left hand is associated with a relative-difference signal
with more values around zero.
The distribution of the values of the relative-differ-
ence signal reflects the balance of co-contraction and
reciprocal activity of the antagonistic muscles. Domi-
nant co-contraction is indicated by a narrow distribu-
tion with the majority of values clustering around zero,
whereas dominant reciprocal activity is indicated by a
wide distribution with only a minority of values clus-
tering around zero, but a majority being close to the
extremes of ±1. The tendency to co-contract the
antagonistic muscles can be assessed by the proportion
of values (and thus the proportion of trial duration)
within a certain range around zero. However, there is
some arbitrariness in defining the boundary of this
range, which corresponds to the relative difference
between myoelectric activity of the antagonistic mus-
cles, which separates the categories ‘‘co-contraction’’
and ‘‘contraction of only one muscle’’. Instead of a
measure of this kind, I used the variance of the rela-
tive-difference signal. The smaller the variance is, the
narrower is the distribution and the stronger is the
tendency to co-contract antagonistic muscles; the lar-
ger the variance is, the wider is the distribution and the
stronger is the tendency toward reciprocal activity.
From Fig. 2, it appears that the modulation of the
relative difference between flexor and extensor activity
is more gradual for the left hand than for the right hand,
for which the changes between extreme values are
rather abrupt. However, the more gradual changes from
the one to the next extreme value of the relative-
difference signal of the left hand are not really smooth,
but superposed by oscillations with a higher frequency
than that of the finger oscillations. Thus the examples
illustrate that the variability of the relative-difference
signal can be more or less closely related to the variation
of the angular position. To capture these differences,
I computed the power spectrum of the relative-differ-
ence signal (after shifting it to a mean of zero) for each
trial (MATLAB function periodogram). The spectra for
the two trials from which the illustrative data of Fig. 2
were taken are shown in Fig. 3 for frequencies up to
15 Hz.

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Exp Brain Res (2007) 178:363–373
The total power of a signal, that is, the integrated
power spectral density, is identical to the variance.
Figure 3 illustrates its decomposition into frequency-
related components, that is, into the variances of the
harmonic oscillations, which are needed to reconstruct
the original relative-difference signal by way of sum-
mation. For each hand the largest component was
clearly at the frequency of the finger oscillations; note
that the frequency was slightly higher for the right hand
than for the left hand. The largest component for the
right hand was larger than the largest component for the
left hand, not only in absolute terms, but also in relative
ones, that is, when expressed as a proportion of total
power. Thus, for the right hand a larger part of the
variation of the relative-difference signal was in the
frequency range of the finger oscillations than for the left
hand. The relative variance in this frequency range was
computed for each trial as the ratio of the power in the
frequency band f ± 1 Hz and the total power, where f is
the frequency of the finger oscillations in that trial (more
precisely, the reciprocal of the median cycle duration).
For each participant the kinematic and myoelectric
measures were averaged across the 20 left-hand trials
and the 20 right-hand trials, respectively. These means
were entered in the statistical analyses.
Results
In a first set of analyses I compared the kinematic and
myoelectric characteristics of finger oscillations with
the dominant and nondominant hand. In these analyses
eventual differences, which resulted from the order of
testing the left and right hand, were taken into account,
but eventual differences between left-handers and
right-handers as well as between men and women were
neglected.
Fig. 3 Power spectra of the
power
relative-difference signals for
1.6
the trials used for the
examples in Fig. 2. For each 1.4
hand there is a clear peak at
the oscillation frequency. This 1.2
peak is smaller both in 1.0
absolute and in relative terms
(relative to total power) for 0.8
the left than for the right hand
0.6
0.4
0.2
0.0
0 2
369
Handedness and gender have been reported to af-
fect manual performance asymmetries (e. g., Peters
and Durding 1979; Carlier et al. 1993). However, the
present experimental design is not fully crossed, and
only a few left-handers were studied. Thus, a compar-
ison between left-handers and right-handers has so
little statistical power that it is essentially meaningless.
Instead a second set of analyses included the 16 right-
handers only. Finally, the effect of gender was ana-
lyzed, but only for right-handers. These additional
analyses are a first step in exploring the generality of
the results across populations known to differ with
respect to manual and cortical asymmetries.
Manual asymmetries
The means of the various kinematic and myoelectric
variables are shown in Table 1. Each of these variables
was subjected to an ANOVA with the within-partici-
pant factor hand and the between-participant factor
order of testing. For the total myoelectric activity the
factor muscle (flexor vs. extensor) was added. The only
significant effects were those of hand; no other main
effects or interactions reached statistical significance.
For the dominant hand the cycle duration was
shorter by about 25 ms, F(1,18) = 27.0, P < 0.01, and it
was less variable, F(1,18) = 18.7, P < 0.01. These dif-
ferences were not accompanied by a difference in
mean amplitude, and there was only a near-significant
trend of the amplitude of the dominant hand to be less
variable than the amplitude of the nondominant hand,
F(1,18) = 3.5, P < 0.10.
Myoelectric activity was significantly stronger in the
dominant than in the nondominant hand, F(1,18) =
16.9, P < 0.01. According to the significant difference
with respect to the variance of the relative-difference
signal, F(1,18) = 4.8, P < 0.05, myoelectric activity was
left hand right hand
power
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
4 6 8 10 12 14 16 0 2 4 6 8 10 12 14 16
frequency [Hz] frequency [Hz]
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370
Table 1 Means of kinematic
and myoelectric variables
observed for finger
oscillations of the dominant
and nondominant hand Cycle duration (ms)
Median
Semi-inter-quartile range
In brackets the means for Amplitude (deg)
right-handers are given Median
a Semi-inter-quartile range
Proportion of total power in Total myoelectric activity (mV s)
the frequency band ±1 Hz Relative-difference signal
around the frequency Total power (variance)
corresponding to the median Relative power at oscillation frequencya
cycle duration of each trial
more reciprocal in the dominant hand, with stronger
co-contractions in the nondominant hand. As indicated
by the relative power in the 2-Hz frequency band
centered at the frequency of finger oscillations, the
reciprocal muscle activity of the dominant hand was
more focussed and less distributed across a range of
frequencies outside the critical band. The manual
asymmetry with respect to relative power was highly
significant, F(1,18) = 37.2, P < 0.01.
Manual asymmetries in right-handers
The analysis of manual asymmetries was repeated for
the sub-sample of 16 right-handers. The means of the
various kinematic and myoelectric variables are also
shown in Table 1 (in brackets). In the ANOVAs again
only significant differences between hands showed up,
but no other main effects or interactions.
In right-handers the cycle duration of the dominant
hand was shorter than the cycle duration of the non-
dominant hand, F(1,14) = 19.8, P < 0.01, and less var-
iable, F(1,14) = 18.6, P < 0.01. Neither for mean
amplitude nor for amplitude variability the difference
between hands approached statistical significance.
Myoelectric activity was significantly stronger in the
dominant hand, F(1,14) = 14.6, P < 0.01, and so was
the variance of the relative-difference signal, F(1,14) =
4.6, P < 0.05, indicating a more reciprocal activity in
the dominant than in the nondominant hand. Finally,
relative power of the relative difference signal in
the 2-Hz band centered at the frequency of finger
oscillations was stronger in the dominant than in the
nondominant hand, F(1,14) = 50.0, P < 0.01.
Modulation of manual asymmetries by gender
In the third set of analyses the manual asymmetries of
male and female right-handers were compared. For the
kinematic variables there was no significant main effect
123
Exp Brain Res (2007) 178:363–373
Hand
Nondominant Dominant
217 (223) 192 (197)
17.1 (17.9) 11.6 (11.8)
32.6 (32.2) 32.9 (32.6)
4.1 (4.1) 3.4 (3.4)
0.295 (0.321) 0.443 (0.466)
0.157 (0.141) 0.204 (0.194)
0.470 (0.437) 0.653 (0.651)
or interaction with gender except for the gender · hand
interaction of the variability of cycle duration,
F(1,12) = 5.2, P < 0.05. For men the mean semi-inter-
quartile ranges for the nondominant and dominant
hand were 15.5 and 12.0 ms, respectively, whereas for
women they were 20.5 and 11.1 ms. (The associated
mean cycle durations were 219 and 192 ms for men,
226 and 201 ms for women.)
Total myoelectric activity exhibited a complex
interaction of gender, order of hands, and hand,
F(1,12) = 7.0, P < 0.05. In general, myoelectric activity
was stronger in the dominant than in the nondominant
hand. Only for women, who were tested first with the
dominant hand, myoelectric activity was stronger in
the nondominant hand, which was tested second. The
variance of the relative-difference signal was signifi-
cantly smaller for women than for men, 0.113 vs 0.212,
F(1,12) = 7.6, P < 0.05. The interaction of gender and
hand only approached statistical significance,
F(1,12) = 4.5, P < 0.10. Nevertheless the manual
asymmetry was present in men only, 0.261 vs 0.163, but
absent in women, 0.113 vs 0.113. For the relative power
in the frequency band around the frequency of finger
oscillations there was no reliable difference between
men and women.
Discussion
Even though the task of the present study was finger
oscillation in the horizontal plane rather than finger
tapping, the kinematic findings are consistent with
those reported in the literature for tapping: perfor-
mance with the dominant hand is faster and tends to be
less variable than with the nondominant hand. The
manual asymmetry with respect to the rate of finger
oscillations was not accompanied by an asymmetry
with respect to amplitude, even though a higher rate is
typically associated with a smaller amplitude (Kay
Exp Brain Res (2007) 178:363–373
et al. 1987; Rosenbaum et al. 1991). At most there was
a tendency of amplitude variability to be smaller in the
dominant than in the nondominant hand.
The replication of typical kinematic asymmetries
shows that the present atypical experimental task does
not result in atypical behavioural findings. The core
findings of the present study, however, are related to
myoelectric activity. Consistent with expectations,
myoelectric activity of the nondominant hand revealed
stronger co-contractions, whereas in the dominant
hand a pattern of reciprocal activity of antagonistic
muscles was more pronounced. The expectations were
based on a generalization of the dynamic-dominance
hypothesis of Sainburg (2002). According to this gen-
eralization, the dominant hand is controlled to exploit
nonmuscular forces, whereas the nondominant hand is
controlled to tame such forces. One way to tame
nonmuscular forces is to increase the visco-elastic
resistance of the joint by way of co-contracting antag-
onistic muscles (Hogan 1984). With sufficiently high
visco-elastic resistance the mass of the physical limb is
tightly coupled to the virtual trajectory defined by the
joint equilibrium position. In contrast, exploitation of
nonmuscular forces, and inertial forces in particular,
requires a low visco-elastic resistance. Reciprocal
bursts of myoelectric activity of antagonistic muscles
are then sufficient to maintain the oscillation of the
mass of the physical limb, the trajectory of which can
deviate considerably from the virtual trajectory.
The distinction between exploitation and taming of
nonmuscular forces does not imply two categorically
different styles of motor control, but rather a contin-
uum. Reciprocal activity of antagonistic muscles is
more prominent in the dominant hand, but it is still
present in the nondominant hand, though superposed
on (relatively) stronger co-contractions. For the dom-
inant hand it has been shown that there are immediate
kinematic consequences of the (relative) timing of the
bursts of reciprocal myoelectric activity on frequency
and amplitude (Heuer and Schulna 2002; Heuer 2006).
Perhaps the stronger variability of the cycle durations
(and perhaps the amplitudes) of the nondominant hand
observed in the present study is related to a stronger
variability of the (relative) timing of the reciprocal
components of myoelectric activity.
Exploitation of nonmuscular forces requires that
these can be anticipated, at least for movements which
are as rapid as the present ones. In line with the dy-
namic-dominance hypothesis, the control of finger
movements of the dominant hand might be based on a
more accurate internal model of the dynamic charac-
teristics of the moving limb than control of finger
movements of the nondominant hand. Such an internal
371
model allows accurate feedforward control, and this
includes accurate temporal placement of bursts of
myoelectric activity. In terms of neural substrates, the
more accurate internal model for the dominant hand
could be related to its more extensive cortical repre-
sentation (Hammond 2002). Consistent with this
hypothesis, Jäncke et al. (2004) found that repetitive
transcranial magnetic stimulation of the left primary
hand motor area eliminated the right-hand superiority
in maximally rapid tapping (all participants were right-
handed in this study).
A higher level of co-contractions is not a unique
characteristic of movements of the nondominant hand
as compared with those of the dominant hand. For
example, it has also been reported for earlier stages of
practicing a tracking task as compared to later stages
(Metz 1970), or for aiming movements of elderly per-
sons as compared to those of younger adults (Darling
et al. 1989; Seidler-Dobrin et al. 1998). Thus it may be
a rather generalized strategy to deal with situations in
which control of dynamic or kinematic transformations
is required without a sufficiently accurate internal
model or with functional impairments due to age-re-
lated changes of the sensori-motor systems.
The present results indicate the possibility that the
manual asymmetry with respect to the level of co-
contractions might be limited to men. Even though the
gender difference was not statistically reliable, there
was a conspicuous absence of the difference in women,
who exhibited a rather high level of co-contractions in
both hands. At present it is not clear whether this
tendency is actually a substantial one, and if it is, what
the reasons for its existence are.
The second major finding of the present study is the
observation that reciprocal myoelectric activity is not
only less pronounced in the nondominant hand than in
the dominant hand, but is also less focussed on the
frequency of the finger oscillations. With a low visco-
elastic resistance, the trajectory of the finger will
approximate a periodic oscillation no matter what the
trajectory of the equilibrium position is. However, with
a high visco-elastic resistance, the trajectory of the
finger will closely match the virtual (equilibrium) tra-
jectory. Therefore, to produce an oscillatory move-
ment, the virtual trajectory should also approximate a
sinusoidal trajectory. Perhaps the generation of such a
smooth trajectory by means of myoelectric activity
with a complex relation to the resultant joint torque is
associated with a noisy character of the relative-dif-
ference signal of the nondominant hand.
Finally I shall consider the possibility that the major
findings of the present study, which concern the variance
(or total power) of the myoelectric relative-difference
123
372
signal and its spectral decomposition, are related to
differences in myoelectric activity or kinematic differ-
ences between the hands. First, consider total myo-
electric activity. The lower the amplitude of myoelectric
signals is, the smaller should be the signal-to-noise ratios
of the signals (cf. Gerdle et al. 1999). This could affect
primarily the relative power of the relative-difference
signal at the frequency of finger oscillations. However,
there is no indication that the manual asymmetry with
respect to total myoelectric activity was related to
manual asymmetries in the other myoelectric variables.
The correlations between manual asymmetries in total
myoelectric activity and in both the variance of the
relative-difference signal and the relative power in the
frequency band around the oscillation frequency were
small and statistically not significant, 0.01 and 0.06,
respectively.
Second, consider the possibility that myoelectric
asymmetries with respect to co-contractions could re-
sult from the kinematic differences, in particular from
the faster oscillations of the dominant than of the
nondominant hand. Again the correlations reveal no
reliable relations between kinematic and myoelectric
asymmetries; the correlations of the intermanual dif-
ference of the variance of the relative-difference signal
with the intermanual differences of cycle duration,
cycle-duration variability, amplitude, and amplitude
variability were –0.01, 0.06, 0.03, and –0.01, respec-
tively. For the relative power of the relative difference
signal in the frequency band around the oscillation
frequency the correlations were 0.10, –0.23, 0.07, and
–0.31, respectively. Even the two negative ones of these
correlations did not reach statistical significance. Thus,
there is no indication that the present findings for
myoelectric activity are a consequence of kinematic
differences, in particular not of the manual asymmetry
with respect to oscillation frequency.
Acknowledgment The research reported in this paper was
supported by grant He 1187/12-1 of the Deutsche Forschungs-
gemeinschaft. I thank Raphael Schulna, Stefan Hohmann, and
Alexander Waschkau for their support in running the experi-
ment, and Alwin Luttmann for stimulating discussions of ways to
analyse EMG signals.
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