J-a -‘, HUMAN

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ELSEVIER Human Movement Science 14 (1995) 217-245

!zfiNT

Coupling strength in tapping a 2 : 3 polyrhythm

C. (Lieke) E. Peper a**, Peter J. Beek a,b, Piet C.W. van Wieringen a
a Institute for Fundamental and Clinical Human Movement Sciences, Faculty of Human Movement
Sciences, Free University, Van der Boechorststraat 9, 1081 BTAmsterdam, The Netherlands
b Center for the Ecological Study of Perception and Action, University of Connecticut, Storrs, USA

Abstract

The effects of tempo and role (fast vs. slow hand) on the reciprocal interaction (strength
of coupling) between two hands tapping a 2: 3 polyrhythm were examined from the
perspective of nonlinear oscillator theory. A measure of the degree of harmonicity was
developed, based on the relative contribution of the tapping frequency to the power
spectrum of the limit cycle phase angle of each individual hand. On the assumption of fixed
coefficients of the dissipative terms in the component oscillators, comparison of unimanual
and bimanual performance with respect to this measure allowed for examination of the
effects of the experimental conditions on the strength of the coupling. Five right-handed
skilled drummers performed the 2: 3 polyrhythm at several tempos and with both hand
arrangements (i.e., either the preferred or the non-preferred hand tapped the faster
cadence). The analysis revealed an inverse relation between tempo and coupling strength,
and a larger influence of the fast hand on the slow hand than vice versa. No differences,
were observed between the two hand arrangements. The theoretical implications of these
results were discussed in relation to similar and dissimilar findings in the literature.

1. Introduction

The coordination of rhythmically moving limbs is characterized by the

interactions between them. The bimanual production of multifrequency
relations provides a case in point. In such performance the hands move at

* Corresponding author. E-mail: c-- e peper@fbw.vu.nl.

0167-9457/95/$09.50 0 1995 Elsevier Science B.V. All rights resewed

SSDI 0167-9457(95)00010-O
218 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

relatively fixed but different frequencies. If the relatively prime ratio of the
movement frequencies contains 1 as numerator or denominator it is re-
ferred to as a ‘simple rhythm’, whereas all other rational ratios constitute
‘polyrhythms’. Although the unimanual subtasks are equally simple,
polyrhythms are performed with more temporal variability than simple
rhythms (e.g., Deutsch, 1983). This difference in accuracy suggests the
presence of interaction between the hands, and the performance of
polyrhythms has been modeled accordingly. Examples of such modeling are
the so-called (integrated) time-keeper models for polyrhythmic perfor-
mance (e.g., Jagacinski et al., 1988; Summers et al., 1993b; Vorberg and
Hambuch, 1984).
However, during the last 15 years or so an alternative view on movement
coordination has been established (Kelso et al., 1981; Kugler et al., 19801,
which holds that the temporal order observed in rhythmic performance is
not prescribed by a motor program but emerges in an autonomous fashion
from the dynamical properties of the system itself. On this view, the
movement system handles its large number of internal degrees of freedom
by functionally organizing itself as a system of nonlinear oscillators which
mutually interact according to a nonlinear coupling function (e.g., Haken et
al., 1985). From mathematical-physical oscillator theory it is known that
such a coupling between oscillators may result in stable behavioral modes
to which the system is attracted. The degree of stability of these modes
depends on the type and the strength of the coupling.
A paradigmatic line of research has been formed by the finger cycling
experiments of Kelso and his colleagues (e.g., Kelso, 1984; Kelso and
Scholz, 1985; Kelso et al., 1986; Kelso and Schbner, 1988). If the movement
frequency of two index fingers cycling in anti-phase (simultaneous contrac-
tion of non-homologous muscle groups) was gradually increased, an abrupt
switch to in-phase coordination (simultaneous contraction of homologous
muscle groups) was observed at some critical frequency. If the fingers
started in the in-phase coordination, however, neither increase nor de-
crease in frequency resulted in such a transition - a phenomenon called
‘hysteresis’. The sudden transition and the hysteresis phenomenon were
captured by a model of two nonlinearly coupled nonlinear oscillators
(henceforth referred to as the HKB-[Haken-Kelso-Bunz-] model), which
described the global behavior of the system in terms of the dynamics of a
single collective variable or ‘order parameter’ (relative phase) and its
dependence on the ‘control parameter’ movement frequency (Haken et al.,
1985).
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 219

An essential assumption of the HKB-model is that linear stiffness, and

hence its dynamical companion frequency, is the only parameter that is
controlled by the nervous system. That is to say, given the composition of
the component oscillators and the functional form of the coupling, and
given a specific initial parametrization, all relevant changes in the space-
time properties of the component oscillators are only due to changes in
frequency. Therefore, dynamical modeling of the abrupt changes in stabil-
ity of coordinative modes, induced by manipulation of the control parame-
ter movement frequency, not only requires a functional description of the
component oscillators (cf. Beek et al., in press-a; Kay et al., 1987) and their
coupling (e.g., Haken et al., 1985; Keith and Rand, 1984; Kopell, 1988;
Rand et al., 19881, but also of the quantitative relation between coupling
strength and movement frequency. Haken et al. (1985) demonstrated that
the phase transition from anti-phase to in-phase coordination was captured
by a potential function for relative phase, which involved two ‘coupling’
parameters that were assumed to decrease differentially with increasing
movement frequency. ’ As a consequence, manipulation of movement fre-
quency involved changes in the ratio between the two parameters, resulting
in changes in the potential function describing the attractor states of the
phase difference between the oscillators.
The general model of coupled oscillators has also been applied success-
fully in situations in which the limbs did not oscillate at the same frequency
but at some multifrequency relation (as in polyrhythmic performance). If
the limbs move at different frequencies, the coordination mode is charac-
terized by the frequency ratio. From oscillator theory it is known that
certain frequency ratios are more stable than others. In general, higher-
order ratios (i.e., ratios composed of large numerators and denominators)
are less stable than lower-order ratios. In line with these theoretical
considerations, several studies on bimanual multifrequency performance
have demonstrated differences in stability between coordination modes: If
the required frequency ratio could not be performed in a stable fashion,
the system was often attracted to a lower-order ratio, indicating that the

’ Haken et al. (198.5) derived the potential function from a system of coupled differential equations
with two coefficients in the coupling function. They demonstrated that the changes in the parameters in
the potential function were associated with these two coupling coefficients in combination with the
drop in amplitude which occurs when movement frequency is increased (due to the Rayleigh term in
the component oscillators). The parameters in the potential function, thus, reflect the degree of
coupling between the oscillators, and hence are sometimes referred to as ‘coupling’ parameters.
220 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

latter ratio was more stable than the former (Kelso and deGuzman, 1988;
Peper et al., in press-b; Treffner and Turvey, 1993). In addition, the
stability of these frequency locks has been demonstrated to depend on
movement frequency. First, attraction to lower-order ratios was more often
observed when the subjects performed at higher than at lower movement
frequencies (Peper et al., in press-b). Second, and relatedly, abrupt transi-
tions to lower-order ratios occurred when movement frequency was gradu-
ally increased (Peper et al., 1991, in press-a; see also Beek et al., 1992a;
Haken et al., in press). Given the assumption that the coefficients of the
nonconservative (dissipative) terms of the component oscillators do not
change when movement frequency is scaled (in conformity to the HKB-
model), these changes in stability must have originated from changes in the
coupling between them.
To understand the abrupt changes in coordination induced by increased
movement frequency, it is of paramount importance to examine the rela-
tion between this control parameter and the coupling parameter(s). The
assumption that the dissipative components of the unit oscillators do not
scale with movement frequency amplifies the focus on the coupling func-
tion in modeling the frequency dependence of pattern stability in rhythmic
performance. Recently, Schmidt et al. (1993) and Sternad et al. (1992) have
successfully addressed this issue with respect to 1: 1 frequency coordination
in swinging hand-held pendulums. They exploited the shifts in the equilib-
rium points of relative phase that were induced by differences in eigenfre-
quency between the component oscillators to examine the relation between
movement frequency and coupling strength. Their analysis was based on
the model proposed by Rand et al. (1988; see also Kopell, 19881, which is in
fact a truncated version of the HKB-model in which the coupling function
consists of a single term with a single parameter (see Fuchs and Kelso,
1994, for a critical note indicating the limitations of this conceptualization).
Experimental results revealed that the coupling parameter was inversely
related to the frequency at which the pendulums were swung.
With respect to multifrequency behavior the problem becomes more
complicated. Haken et al. (in press) demonstrated that, in order to account
for the relatively large number of observed stable ratios and for the
frequency-induced transitions between them, a coupling function with
many terms and associated parameters is required. Consequently, the
empirical examination of coupling strength and changes therein when a
control parameter is scaled is not as straightforward as in 1: 1 frequency
locking. In the next section, we present a general procedure for examining
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 221

the relative magnitude of coupling strength in rhythmic performance. It

may be applied whenever the coupling function involves multiple terms and
parameters and even in case the precise coupling function is unknown.
Rather than examining the effects of the coupling on the value of the
collective variable (cf. Schmidt et al., 1993; Sternad et al., 19921, the
proposed method measures the effects of the coupling on the movement
patterns of the individual limbs. Therefore, it readily allows for examina-
tion of asymmetries in the coupling between the oscillators. Although the
coupling function in the HKB-model is symmetrical (the oscillators interact
via identical coupling terms), recent findings suggest asymmetries in the
interaction between the limbs. Byblow et al. (1994) and Byblow and
Goodman (1994) have reported differences in performance between the
right (preferred) and left (non-preferred) hand during bimanual frequency-
locked behavior. They suggested that this asymmetry resulted from differ-
ences between the hands with respect to the degree to which coupling
strength scales with movement frequency (see also Carson et al., 1993). In
testing the performance of multifrequency ratios in skilled drummers,
Peper et al. (in press-b) created an asymmetrical situation in which one
hand had a larger influence on the other hand than vice versa. Their
subjects were able to perform ratios of relatively high order as long as the
‘forcing’ hand tapped the faster frequency. When this forcing hand moved
slower than the other hand, however, significantly more transitions to
lower-order ratios were observed. Contrary to the findings of Byblow et al.
(1994) and Byblow and Goodman (19941, performance was similar when
either the preferred (right) or the non-preferred (left) hand functioned as
‘forcer’. These results suggested that multifrequency performance is char-
acterized by an asymmetrical coupling in that the fast hand has a larger
influence on the slow hand than vice versa, which is in agreement with
previous observations for polyrhythmic performance (Peters and Schwartz,
1989; Summers et al., 1993a,b). Within the framework of the general model
of coupled oscillators, the suggested asymmetries in the coupling between
the hands provide plausible explanations for the observed effects. However,
the postulated differences between the hands with respect to the coupling
influences that they exert onto each other have not yet been empirically
demonstrated. Because the method introduced in the next section allows
for quantification of the strength of interaction between the limbs on the
basis of examination of the effects on the individual movement patterns,
asymmetries in coupling strength can be examined. In the present experi-
ment, the suggested asymmetries resulting from differences between the
222 C.E. Peper et al. /Human Movement Science I4 (1995) 217-245

hands as well as those resulting from differences between the functional

roles of the hands (fast vs. slow hand) were thus studied.

2. A method to determine the relative coupling strength

In line with the research reviewed in the introduction, we model the

oscillating limbs as a system of nonlinearly coupled nonlinear oscillators.
Following Haken et al. (1985), the functional form of the component
oscillators and the mutual coupling is assumed to be fixed. In general terms
the differential equations read:

and
&+qx,,&) +g(-Q) =I,,,
where xi and x2 refer to the positions in time attained by the two
oscillators, using the dot notation to indicate time derivatives. I,, and I,,
represent the coupling influences of the two oscillators onto each other.
The dissipative terms in h(x,, ii> are assumed to be weakly nonlinear,
involving both positive and negative damping: The component oscillators
are self-sustaining and their long-term behavior is, thus, independent of the
initial conditions. When represented in the phase plane (i.e., i plotted
against x), the trajectory is attracted to a closed curve: the limit cycle. For
weakly nonlinear dissipation this pattern can be an almost perfect circle
indicating near-harmonic behavior. In the model, the two oscillators inter-
act through the coupling functions I. The oscillators ‘force’ each other,
which results in increased nonlinearity (relaxation) in the movement pat-
terns. This effect of coupling may be exploited to examine the degree of
interaction between the limbs in movement coordination by comparing the
extent to which the behavior is harmonic under unimanual and bimanual
conditions. The present analysis, therefore, concentrates on the influence
of experimental manipulations on the degree to which the movements are
harmonic. The degree of harmonicity is operationalized as the relative
contribution of the tapping frequency to the power spectrum of the
evolving phase in the oscillatory pattern. Because the behavior of a limit
cycle oscillator is essentially captured in its phase plane, defined by the
orthogonal coordinates
x=rcos+, i=rsin4, (2a,b)
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 223

we focus on the limit cycle angle variable (4). In the context of rhythmic
tapping it is convenient to take this measure modula 27r, so that the
distortions in the time series caused by missed or irregular taps have less
influence on the power spectra than they would have in a continuous
measure of 4. The spectrum of the resulting saw-tooth-like evolution of $
shows a large peak at the required frequency and additional smaller peaks
at its higher harmonics. The effects of frequency and nonlinearity on the
power spectra of a limit cycle oscillator are illustrated in Fig. 1, which
represents the results of the proposed spectral analysis on numerical
simulations of the Van der Pol oscillator. ’ Inspection of these power
spectra reveals that the peak at the required frequency decreases for larger
degrees of nonlinearity, both in absolute measures and relative to the other
spectral peaks (compare Figs. 1A and lC, and Figs. 1B and 1D). This effect
is less pronounced if the system oscillates at higher frequencies (compare
Figs. 1C and 1D). The relative contribution of the peak at the required
frequency (taking a peak width of + 10% of Trequired)to the total power of
the spectrum indicates a similar effect (Fig. 1A: 60.8%; B: 60.6%; C:
57.4%; D: 60.0%). In sum, the contribution of the peak at the required
frequency relative to the spectral peaks at the higher harmonics is related
to the relaxation component in the oscillatory pattern. These results are in
line with Fuchs et al. (submitted), who demonstrated, both numerically and
analytically, that if the phase trajectoy of an oscillator is not circular (e.g.,
due to nonlinearities) complicated oscillating behavior is observed in 4
(when determined on the basis of Eqs. (2a,b)).
Given our assumption of a fixed dissipative structure of the component
oscillators, the contribution of the actual tapping frequency to the power
spectrum, when considered relative to the contribution of its higher har-
monics, reflects the degree to which the oscillatory pattern is influenced by
the other oscillator. A larger degree of relaxation (larger contribution of
the higher harmonics) is associated with stronger coupling. Therefore,
comparison of bimanual and unimanual performance with regard to the
relative contributions of the actual tapping frequency to the power spectra

* Kay et al. (1987) successfully modeled oscillatory hand movements as hybrid oscillators, involving
both a Van der Pol and a Rayleigh dissipative term. The degree to which the behavior is nonlinear
depends on the relative weights of these two terms. Because for nearly harmonic behavior the gain
parameter associated with the Rayleigh term is small compared to that of the Van der Pol term (Kay et
al., 1987), we chose, for reasons of clarity, to illustrate the effects of increased nonlinearity by
considering the behavior of an oscillator with only a Van der Pol term.
224 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

time(s) lime (s)

,,u--_
0 I? 3 4 5 6 0 3 6 9 12 15 18

frequency (Hz) frequency (Hz)

I_...i_._L_
3 6 9 1?
1

IS 18

frequency (Hz)
Fig. 1. Limit cycle phase (mod 2~) and power spectrum thereof for a numerically simulated Van der
Pol oscillator (a + ai + ytix2 + 02x = 0; fourth-order Runge Kutta integration method). Time step =
0.01 s. Initial conditions: x0 = 1.15 (cm), i, = 0 km/s). Parameters: Panel A: 1y= -0.3 (Hz), y = 0.9
(Hz/cm), w = 6.28 (rad/s); Panel B: Q = -0.3 (Hz), y = 0.9 (Hz/cm), w = 18.85 (rad/s); Panel C:
a = - 4.8 (Hz), y = 14.4 (Hz/cm), w = 6.28 (rad/s); Panel D: a = -4.8 (Hz), y = 14.4 (Hz/cm),
w = 18.85 (rad/s). Increase in y (together with a) implies increased nonlinearity (relaxation); increase
in 0 reflects increase in movement frequency f (w = 2pf).

under different experimental conditions provides a measure for the relative

magnitude of the coupling strength in these conditions. On the basis of
these considerations, hypotheses with respect to the effects of tempo and
role (fast hand vs. slow hand) on the interaction between the hands in
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 225

polyrhythmic tapping were formulated. The present experiment was de-

signed to test these hypotheses. Moreover, we examined whether perfor-
mance was affected when the roles of the hands were exchanged between
the preferred (right) and the non-preferred (left) hand.

3. Experiment

In this experiment three hypotheses were examined using the proposed

analysis based on changes in relaxation in the movement patterns: (i) the
relaxation component in bimanual performance is larger than in unimanual
performance (reflecting the coupling between the hands); (ii) if movement
frequency is scaled down, the degree of relaxation increases more strongly
in bimanual than in unimanual tapping (reflecting an inverse relation
between movement frequency and coupling strength); (iii) the relaxation
component in the movement pattern of the slow hand is larger than in that
of the fast hand (reflecting an asymmetrical coupling between the hands).
To test these hypotheses the subjects were invited to tap a 2: 3 polyrhythm
at six different tempos (overall cycle frequencies). Each subject performed
the bimanual task under two hand arrangements, in which either the right
(preferred) or the left (non-preferred) hand performed the faster cadence,
to allow for examination of possible effects of hand preference (cf. e.g.,
Byblow et al., 1994; Byblow and Goodman, 1994). In addition, all compo-
nent frequencies were also performed, with both hands individually, in
unimanual control trials.

3.1. Method

Subjects
Five skilled drummers participated in the experiment (mean age: 28.6
years, range: 20-34). In terms of the way in which they were used to
drumming, they were all right handed (playing the hi-hat with the right
hand and left foot, the snare with the left hand, and the base with the right
foot). They were paid for their services.

Experimental set-up
The hand movements were measured with a 2D-Selspot system which
recorded the position of two Light Emitting Diodes (LEDs) positioned on
the tips of the middle fingers (sample frequency 312 Hz). A micro-com-
226 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

puter (IBM PS/2 40 SX> controlled the Selspot system and the stimulus
device which produced auditory stimuli. The accuracy of the stimulus onset
was f 1.5 ms. Rhythmic stimulus trains (duration 35 s), consisting of 50 ms
beeps, were presented through the two channels of a headphone (Senn-
heiser HD 520 II). The pitch of the tones was different for the two
channels (right: 440 Hz; left: 200 Hz). 3 The taps were performed on a
low-resonance marble tabletop surface.

Procedure
The subject was seated in upright position (wearing the headphone) and
was instructed to tap with his hands on the tabletop (rotation around the
wrist) while the lower arms were resting on its surface. The right hand had
to synchronize to the stimuli on the right channel of the headphone, the
left hand to those on the left channel. The experiment consisted of two
parts. In the first part the subject performed unimanually along with a
single stimulus train. In the unimanual trials the required frequencies were
presented for one of the hands (blocked; left-right order counterbalanced
over subjects). The eleven frequencies that would be used in the bimanual
trials in Part 2 (interbeep intervals: 300, 400, 450, 500, 600, 700, 750, 800,
900, 1050, and 1200 ms) were tested in blocks of two repetitions, which
were presented in random order.
In the second part two stimulus trains were presented, the frequencies of
which related as 2: 3. This frequency ratio was presented at six cycle
frequencies. The interbeep interval of the faster stimulus ranged from 300
to 800 ms, in 100 ms steps. The interbeep intervals of the slower stimulus
train were chosen such that the 2: 3 frequency ratio was attained, resulting
in interbeep intervals ranging from 450 to 1200 ms (in 150 ms steps). Each
stimulus train could be presented on either the left or the right channel of
the headphone. These situations were tested blockwise (left-right order
counterbalanced over subjects). Within such a ‘hand’ condition four blocked
repetitions of each frequency condition were presented at random. The
first trial in such a frequency block functioned as practice trail. Thus, 36
experimental bimanual trials were conducted in total. This second part of

3 The pitches were, thus, noticably different. However, this difference was small and the pitches were
comparable to those that were selected by Jagacinski et al. (1988) to support the perception of an
integrated pattern, which has been demonstrated to have a benificial effect on the accuracy of
performance of polyrhythms (Jagacinski et al., 1988; Knapp et al., 1985).
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 227

the experiment was preceded by three practice trials‘in which the interbeep
interval of the faster stimulus train was 600 ms.

Analysis
The Selspot data were converted to Cartesian coordinates (using Direct
Linear Transformation, cf. Miller et al., 1980; Shapiro, 1978) and filtered
(second-order recursive Butter-worth, applied back and forth, cut-off 25
Hz). To double the number of tapping cycles for which the power spectra
would be derived (see below), the time series were resampled at 150 Hz.
The frequency of oscillation was determined for each tapping cycle, using a
peak picking algorithm to determine the moments in time of peak exten-
sion (7’pE) for both hands. From these TPEs the angular frequency for
every nth cycle was calculated as

f,= 27T
TPE, +, - TPE, *

For each cycle the center of oscillation (C) was determined using the
LED-positions attained at peak extension (PPE) and when the hand
contacted the table (no extension: PNE), defining every nth PPE to
succeed the nth PNE
PNE, + PNE, +1
2

In other words, the center of each tapping cycle was determined halfway
the extremes of the movement. The amplitude of the movement (xi) was
defined as the displacement of the position value at sample i in the nth
cycle (i.e., between PNE, and PNEn+l) from the corresponding C,. The
limit cycle phase angle at sample i ($J was defined as

&i = arctans (mod 2~), (5)

1

where ii is the velocity at sample i. The phase plane representation was

scaled (‘normalized’) by multiplication of xi with the mean angular fre-
quency obtained for the correspondig cycle (f,): xi* = xi f, (see Beek and
Beek, 1988). It was convenient to define the phase at PPE, as 4 = 0, and
at PNE,,,, as 6 = W. Fig. 2 presents two examples of the resulting phase
evolution over time. The power spectra of 4 were determined for 4096
228 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

FAST HAND (3.3 Hz)

FAST HAND (1.25Hz)

001 ” i -.r
00 05 10 I5 2” 25 30

tmr (S)

Fig. 2. Typical phase evolutions as obtained for the right hand of Subject C during unimanual
performance of intertap intervals 300 ms (upper panel) and 800 ms (lower panel).

samples (27.4 s, omitting the first 5 s and the last 2.6 s of the time series)
using Fast Fourier Transform. To reduce the error, the spectra obtained
for trials from the same condition were averaged for each subject. All
power spectra revealed marked peaks at the required frequency and higher
harmonics thereof (see Fig. 3). The contribution of the most prominent
spectral peaks was determined by calculating the power within the range of
f 10% of T, where T represents the period (inverse of frequency) for
which a peak was observed. The peaks that contributed at least 5% of the
power accounted for by the largest peak were selected for further analysis.
The statistical analysis concentrated on the percentage of the power
summed over the selected peaks that was accounted for by the required
frequency’ Prequired p~ak/~all peaks X 100%. Larger percentages reveal stronger
harmonicity (which is equivalent to a smaller relaxation component).
 C.E. Peper et al. /Human Movement Science 14 (I 99.5) 217-245 229

UNIMANLJAL BIMANUAL
3.33Hz 2.22Hz 3.33Hz 2.22Hz

25. 25 257 251

20: 1 20 20. 201

00
0 2 4 6 8 1012
frequency (Hz)
05:
001i ‘iI1
0 2 4 6
frequency (Hz)
l,

8 1012 0 2 4 6 8 IO 12
frequency (Hz)
0 2 4 6 8 IO I2
frequency (Hz)
UNIMANUAL BIMANUAL
1.25Hz 0.83Hz 1.25Hz 0.83Hz

12. 127 I? I2
I

" 2 4 6 0 2 4 6 0 2 4 6 0 2 4 6
frequency (Hz) frequency (Hz) frequency (Hz) frequency (Hz)

Fig. 3. Typical power spectra as obtained for Subject C. The averaged spectra for the fast (right) and
the slow (left) hand for two tempo conditions (300 ms: upper panels; 800 ms: lower panels) during
unimanual as well as bimanual performance are presented.

3.2. Results

Spectral analysis
The selected spectral peaks accounted for 71.3% (SD = 11.2%) of the
total power in the spectra, averaged over subjects and conditions. The
effects of the experimental conditions on the percentage of power ex-
plained by the required frequency were examined in a 2 x 2 X 2 X 6 Analy-
sis of Variance (ANOVA) with repeated measures, testing the influence of
the factors number of hands (unimanual vs. bimanual), role (fast vs. slow
hand), hand (right vs. left hand) and tempo (6 levels) on the relative degree
of harmonicity (Prequired peak/Pa,, peaksX 100%). The effect of role was
significant, F(1,4) = 154.80, p < 0.0005, revealing that the degree of har-
monicity was larger in case the hand performed the higher frequency (fast
hand: 50.5%; slow hand: 36.5%). The main effect of tempo was also
significant, F(5,20) = 17.26, p < 0.0001. For the range of intertap condi-
tions from 300 to 800 ms (specified for the fast hand) the percentages of
power accounted for by the required frequency decreased as follows:
58.1%, 47.4%, 41.4%, 38.9%, 37.6%, and 37.6%. Post-hoc analysis (New-
man-Keuls, p < 0.05) revealed significant differences between the 300 ms
230 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

04 :‘: :‘:‘: : :‘i’: :‘:‘I

0 100 200 300 300 500 600 700 800 900 1000 1100 ,200

Intellap lllterval (Ills)

Fig. 4. The relative power contribution of the required frequency Charmonicity’), averaged over
subjects, as a function of intertap interval. Note that the ranges of intertap intervals differ for the fast
and the slow hand.

situation and all other tempo conditions, and between the 400 ms condition
and the four slower situations. Number of hands showed a significant
interaction with role (unimanual: fast hand: 50.8%, slow hand: 43.4%;
bimanual: fast hand: 50.4%, slow hand: 29.5%), F(1,4) = 19.53, p < 0.05.
Post-hoc comparison (Newman-Keuls, p < 0.05) revealed a significantly
smaller degree of harmonicity for the slow hand in the bimanual situation
than in the other Number of Hands X Role combinations. The interaction
between tempo and number of hands tended toward significance, F(5,20)
= 2.71, p < 0.1, suggesting that the effect of tempo was mainly due to the
decrease in harmonicity in the bimanual situation (see also Fig. 4).
Because the fast and the slow hand performed at different movement
frequencies, the obtained main effect of role may (partly) reflect the effect
of tempo rather than a genuine effect of the difference between the roles
that the hands performed. To examine this possibility, two additional
repeated-measures ANOVAs were conducted. These ANOVAs were simi-
lar to the previous one, except for the fact that role (fast vs. slow hand) was
excluded as a factor and that the data were subdivided into the two role
classes, together with the corresponding unimanual control trials. For the
slow hand, tempo had a significant effect on the degree of harmonicity,
F(5,20) = 13.15, p < 0.0001. Harmonicity declined over the range of inter-
tap intervals (450 to 1200 ms) in the following way: 50.0%, 39.3%, 35.5%,
33.3%, 30.6%, and 30.0%. Post-hoc analysis (Newman-Keuls, p < 0.05)
showed significant differences between the 450 ms intertap interval (300 ms
condition) and the other 5 tempos, and between the 600 ms interval (400
ms condition) and the two slowest tempos. The effect of number of hands
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 231

tended toward significance, F(1,4) = 4.95, p < 0.1: In the unimanual situa-
tion the movement patterns tended to be more harmonic than in bimanual
performance (43.4% and 29.5%, respectively). In addition, the Number of
Hands X Tempo interaction was significant, F(5,20) = 3.28, p < 0.05. Post-
hoc comparison (Newman-Keuls, p < 0.05) revealed that the tempo effect
was only observed for bimanual performance (see Fig. 4). The second
ANOVA revealed that also when the hand tapped the faster cadence (or
performed unimanually) the degree of harmonicity was significantly influ-
enced by tempo, F(5,20) = 7.52, p < 0.0005. The percentages obtained for
the range of intertap intervals from 300 to 800 were: 66.1%, 55.4%, 47.3%,
44.6%, 44.6%, and 45.1%, respectively. Post-hoc analysis (Newman-Keuls,
p < 0.05) revealed that only the 300 ms condition resulted in a significantly
stronger degree of harmonicity than any of the other conditions.

FAST HAND (2 Hz)

0 I 2 3 4 5 6 7 8

frequency (Hz)

SLOW HAND (1.33 Hz)

06

0 I 2 3 4 5 6 7 8

freqoe%lcy(Hz)
Fig. 5. Examples of additional peaks in the power spectra as observed for Subject E in the 500 ms
conditions. The arrows indicate the additional peaks that were selected for analysis.
232 C.E. Peper et al. /Human Movement Science I4 (1995) 217-245

Detailed inspection of the power spectra revealed a difference between

the unimanual and bimanual conditions. Whereas in the unimanual trials
peaks were only observed at the required frequency and its higher harmon-
ics, bimanual tapping sometimes resulted in a number of small additional
peaks. We could categorize all observed additional peaks into two classes.
In most cases the extra peaks were observed for the slow hand at one half
of the tapping frequency of this hand (i.e., at 0.5 ~~~~~~~or at odd integer
multiples thereof. In a smaller number of cases an extra peak was observed
in the spectrum of the fast hand. These peaks were at frequencies that
coincided with frequencies obtained for the slow hand (including the ones
previously referred to). In both cases these extra peaks were relatively

Subject A Subject B

450 600 750 900 1050 1200 450 600 750 900 lOSO ,200

KItertapInterval (Ins) Interrap interval (ms)

Subject D Subject E

450 600 750 900 ,050 1200 450 600 750 900 1050 1200

Intertap mterval (ms) Intertap Interval (ms)

Fig. 6. The percentage of power accounted for by the additional peaks in the spectrum of the slow
hand, presented as a function of intertap interval for four subjects separately (black: right hand; white:
left hand).
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 233

Table 1
Percentage of peak power explained by ‘additional peaks’ in the spectra obtained for the fast hand. See
text for details
Intertap interval (ms.1 Subject B Subject E
Right hand Left hand Right hand Left hand
300 0 7.2 0 0
400 0 0 0 0
500 0 4.4 7.0 0
600 0 3.8 6.3 14.2
700 0 0 4.1 10.3
800 23.9 5.7 4.1 10.3

small (cf. Fig. 5). The power contributed by these peaks and the conditions
under which they were observed differed across subjects. Subject C did not
show a single additional spectral peak in any of the conditions. Considering
the first type of extra peaks (at odd integer multiples of 0.5 xf,,,,; see Fig.
6), we found that for Subject A the percentage of total power accounted for
by these additional spectral peaks was very small. For this subject and for
Subject B they occurred predominantly in trials in which the right hand was
the fast hand: The extra peaks were then observed for the slower left hand.
For Subject D the opposite effect was obtained. In Subject E additional
spectral peaks were observed in both bimanual hand conditions, although
the contribution to the power spectrum was slightly higher if the right hand
performed the slower cadence. The second type of additional peaks (in the
spectra of the fast hand) were exclusively observed for two subjects. Table 1
shows that these extra peaks occurred predominantly in the slower fre-
quency conditions.

Temporal partitioning of the tapping cycle

The spectral analysis revealed systematic changes in the degree of
relaxation in the tapping movements. In addition, it revealed that even in
unimanual tapping the movements were not altogether harmonic. Tapping
the hands on a rigid surface resulted in temporal patterns involving a flight
period (Tflight) and a period following the actual tap during which the hand
remained on the surface (Tsurface 1. Increases in relaxation may be related to
increases in Tsurface.In evaluating the results of the spectral analysis it is,
therefore, important to examine the effects of the experimental conditions
on the temporal partitioning of the tapping movements. For this reason we
determined, per subject, the mean values of Toight and Turface for each
234 C.E. Peper et al. /Human Mocwment Science 14 (1995) 217-245

0 ‘:‘:‘i :‘:‘:‘/‘i’:‘i’:‘I

0 100 200 300 400 500 600 700 800 900 1000 1100 I200

. *.: .
d
0

o-e-, I I / : ‘,

0 100 200 300 400 500 600 700 800 900 1000 1100 1200

1nrertapInterval (ms)
Fig. 7. Absolute flight period (upper panel) and relative flight period (lower panel), both averaged over
subjects, as a function of intertap interval. Note that the ranges of intertap intervals differ for the fast
and the slow hand.

condition. As is evident from Fig. 7 (upper panel), Tflight increased with

increasing intertap interval. In additional analyses we addressed the ques-
tion as to whether the temporal partitioning of the tapping cycle could
simply be explained by a model of proportional duration, involving a fixed
ratio between the flight period and the mean intertap period 4
(Tflight/Tintertap~ where ‘intertap = Lrface + Tflight). Fig. 7 (lower panel) repre-
sents the values of this variable averaged over the subjects. To examine the
effect of the experimental conditions on Tflight/Tintertapa set of ANOVAs
was conducted.

4 We reserve the term ‘intertap interval’ to indicate the tempo conditions. whereas ‘intertap period’
refers to the actually observed period between the taps.
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 235

A 2 (number of hands) X 2 (hand) X 2 (role) X 6 (tempo) ANOVA with

repeated measures was performed on the percentage of the intertap period
accounted for by the flight period: T”ight/Tintertapx 100%. The effect of role
was significant (fast hand: 65.8%; slow hand: 52.6%), F(1,4) = 36.9, p <
0.01. Tempo also caused a significant main effect, F&20) = 15.2, p <
0.0001. Over the range of intertap intervals from 300 to 800 ms (as defined
for the fast hand) the proportion of the intertap period accounted for by
the flight period was 74.6%, 64.4%, 58.5%, 54.0%, 51.7%, and 52.1%,
respectively. Post-hoc comparison (Newman-Keuls, p < 0.05) revealed that
the 300 ms condition resulted in larger values than all other conditions and
that the percentages obtained in the 400 ms condition were larger than
those in the 600, 700, and 800 ms conditions. Role and tempo showed a
significant interaction, F(5,20) = 2.9, p < 0.05. Post-hoc analysis
(Newman-Keuls, p < 0.05) revealed that this interaction was due to a
stronger effect of tempo for the fast hand than for the slow hand. In
addition the interaction between number of hands and role tended toward
significance (unimanual: fast hand: 66.1%, slow hand: 58.1%; bimanual:
fast hand: 65.5%, slow hand: 47.2%), F(1,4) = 6.5, p < 0.1.
Additional ANOVAs were performed, for the slow and the fast hand
separately, to dissociate the role and tempo effects. The 2 X 2 X 6 re-
peated-measures ANOVA performed on the data obtained for the slow
hand during bimanual performance and for the corresponding unimanual
conditions, revealed a significant effect of tempo, F(5,20) = 10.4, p <
0.0001. For the range of intertap intervals (450 to 1200 ms) the obtained
percentages were: 64.0%, 58.4%, 51.6%, 46.5%, 46.8%, and 48.4%, respec-
tively. Post-hoc analysis (Newman-Keuls, p < 0.05) revealed that both the
450 and the 600 ms intertap intervals (300 and 400 ms conditions, as
defined for the fast hand) resulted in larger values than the four slower
conditions. The effect of number of hands tended toward significance
(unimanual: 58.1%, bimanual: 47.2%), F(1,4) = 6.2, p < 0.1. A similar
ANOVA conducted on the percentages obtained for the fast hand during
bimanual performance and for the corresponding unimanual conditions,
revealed a main effect of tempo, F(5,20) = 12.5, p < 0.0001. For the range
of intertap intervals (300 to 800 ms) the relative flight periods decreased as
follows: g5.3%, 70.5%, 65.4%, 61.5%, 56.5%, and 55.8%. Post-hoc analysis
(Newman-Keuls, p < 0.05) revealed that the 300 ms intertap interval re-
sulted in larger values than all other conditions and that the percentages
obtained in the 400 ms intertap interval were larger than those in the 700
and 800 ms intervals. In addition. the interaction between number of hands
236 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

and tempo turned out to be significant (cf. Fig. 7), F(5,20) = 2.8, p < 0.5.
Post-hoc analysis (Newman-Keuls, p < 0.05)revealed that the tempo effect
was stronger if the subject performed bimanually.

4. Discussion

In the dynamical modeling of rhythmic movements, the interaction

between the limbs is captured by a nonlinear coupling function between
nonlinear oscillators. Assuming that the parameters of the dissipative terms
of the unit oscillators are fixed (i.e., independent of frequency and other
task conditions), changes in coordination result from changes in the cou-
pling parameter(s). To investigate the strength of interaction we capitalized
on the influence of the coupling on the kinematics of the oscillations:
Given the model assumptions, the degree of relaxation in the movement
pattern obtained for one of the oscillators was associated with the coupling
influence of the other oscillator. To examine the changes in coupling
strength induced by experimental manipulations, we developed an index of
the relative degree of relaxation. This measure was operationalized as the
contribution of the spectral peak corresponding to the actual tapping
frequency relative to the other peaks in the power spectra of the evolving
limit cycle phase, as obtained for the movement patterns of the two hands
separately. The proposed analysis may be applied when the exact coupling
function is not known or when it involves multiple terms and parameters,
rendering a more direct investigation impossible or troublesome. We ap-
plied this index of the relative degree of relaxation to movement patterns
observed in tapping a 2: 3 polyrhythm. The analysis was used to determine
the effects of tempo, role (fast vs. slow hand), and hand (left vs. right hand)
on the degree of relaxation in the tapping movements. Unimanual control
trials were included in the analysis to test the hypothesis that changes in
relaxation were indeed caused by the interactions between the hands and
not by changes in the oscillator characteristics of the individual hands.
It is important to realize that the changes in harmonicity can only be
attributed to changes in coupling strength to the extent that the model
assumptions are valid. Recent work by Beek and colleagues indicated that
the assumption that linear stiffness is the only parameter controlled by the
nervous system is an oversimplification. Detailed examination of movement
kinematics in rhythmic tasks revealed that the control of rhythmic move-
ment also involves nonlinear stiffness and friction terms (Beek et al., 1992b,
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 237

in press-b) which may scale with movement frequency (Beek et al., in

press-a). For this reason it is important to include unimanual control data
in analyses that rely on the simplifying assumption of fixed dissipative
terms. In that way the effects of frequency on the component oscillators
can be examined and can be compared to the changes observed in biman-
ual performance. The differences thus obtained are to be understood as a
result of the coupling between the limbs. In this respect we note that
although the effect of tempo, in the present data set, was strongest in
bimanual tapping, the fact that also for unimanual performance a tempo
effect was obtained might indicate that manipulation of movement fre-
quency resulted in changes in the dissipative characteristics of the unit
oscillators. However, two considerations are in order here. First, the
change in harmonicity may be (partly) due to the fact that, in case the
dissipative terms remain unchanged, increase in movement frequency is
associated with decrease in relaxation (compare Fig. 1C and 1D). Second,
because coupling between oscillators is by no means limited to mechani-
cally connected systems (e.g., Schmidt et al., 1990; Wimmers et al., 1992)
the observed effect may (in part) be the result of the unidirectional
coupling influence of the periodic pacing signal. Be that as it may, the
difference between the bimanual and unimanual situation revealed that, as
hypothesized, coupling strength increased with decreasing movement fre-
quency. In what follows we will concentrate on the effects of the experi-
mental manipulations on the strength of interaction between the hands, as
evident from comparisons between unimanual and bimanual performance.
As mentioned above, increase in movement frequency resulted in a
decrease in the degree of relaxation in the movement patterns. This effect
tended to be stronger in the bimanual situation. In addition, the degree of
relaxation was larger for the slow hand than for the fast hand. The
interaction effect between number of hands and role revealed that this
difference between the fast and the slow hand was not simply an artifact of
the difference between the movement frequencies at which the hands
tapped, but was indeed caused by the difference in role during bimanual
tapping. Additional ANOVAs performed for the slow and the fast hand (as
defined for bimanual performance) separately, revealed even more clearly
that the. differences in role resulted in an asymmetry in the mutual
coupling. For the fast hand, the unimanual and bimanual conditions did
not result in significant differences with respect to the relative degree of
relaxation or its depedence on tempo, indicating that this hand was not
noticably influenced by the coupling to the slow hand. However, the
238 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

performance of the slow hand was less harmonic during bimanual tapping
than in the unimanual control trials. In addition, the influence of move-
ment frequency on the movement patterns of this hand was significantly
stronger in the bimanual situation, revealing that the obtained tempo effect
in overall bimanual performance was caused predominantly by the influ-
ence of the fast hand on the slow hand.
Given our assumptions, changes in relaxation reflect changes in the
strength of coupling. The converging results, therefore, support all three
hypotheses: (i) In the bimanual situation relaxation was larger, reflecting
the interaction between the hands; (ii) The effect of tempo, which was
strongest in the power spectra obtained for the slow hand in bimanual
tapping, revealed that coupling strength decreased as movement frequency
increased; and (iii) The coupling between the hands was asymmetrical, as
was evident from the influence of the role that was assigned to the hand:
Whereas the slow hand was influenced by the fast hand, the latter was not
noticably influenced by the former. No differences were observed between
the left and right hand, suggesting no asymmetry in coupling due to the
(right-) hand preference of our subjects. The support of hypothesis (i)
corroborates the notion that the interaction between the limbs in coordi-
nated polyrhythmic behavior may be understood in terms of a coupling
function between coupled oscillators. The support of hypothesis (ii) is in
line with earlier studies on the relation between movement frequency and
coupling strength (Schmidt et al., 1993; Sternad et al., 1992). Whereas
these previous studies examined the effects of movement frequency in
terms of the collective variable relative phase, the present analysis of
coupling strength was based on the kinematics of the movement patterns of
the individual limbs. The fact that the same conclusion was reached along
different lines of analysis certainly adds to the credibility of hypothesis (ii).
We discuss the implications of the evidence for hypothesis (iii) in more
detail in the next section.

4.1. Asymmetrical coupling

The present results indicate that the asymmetry between the fast and the
slow hand in bimanual polyrhythmic performance is rather strong. The
degree of harmonicity in the movement patterns of the hand tapping the
faster frequency during bimanual performance was not significantly differ-
ent from its value during unimanual performance, whereas such a differ-
ence did exist for the slow hand. In addition, the tempo effect was most
 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245 239

pronounced for the slow hand during bimanual tapping. In combination,

these results suggested an almost unidirectional coupling in that the fast
hand strongly influenced the slow hand without being itself much influ-
enced by this other hand. For this reason, the often used sine circle map, a
difference equation modeling the behavior of a unidirectionally forced
oscillator (see below), seems to be more appropriate than one might expect
intuitively. As discussed by Peper et al. (in press-b) the asymmetry in the
coupling is functional for the performance of polyrhythms. When the
forcing oscillator moves at a higher frequency than the forced oscillator,
more frequency relations can be attained. A similar effect was demon-
strated in numerical studies on chemical oscillators (M. Dolnik, personal
communication, November 18, 1993 and September 12, 1994; Kai and
Tomita, 1979; Schreiber et al., 1988).
The evidence for an asymmetrical coupling is in accordance with the
results obtained in earlier studies on polyrhythmic tapping which showed
that the timing of the slow hand depended more on the timing of the fast
hand than vice versa (Peters and Schwartz, 1989; Summers and Kennedy,
1992; Summers et al., 1993a,b). This effect was found to be most pro-
nounced in experienced musicians, suggesting that the identified kind of
dependence reflects the skilled way to perform polyrhythms. In this respect
it may be noted that the subjects in the present experiment were skilled
drummers.
Previous studies have revealed manual asymmetries in rhythmic biman-
ual performance: If tapping a steady beat with one hand is combined with
tapping a rhythmic pattern or tapping as fast as possible with the other
hand, right-handed subjects performed most accurately if the left hand
performed the steady beat (Ibbotson and Morton, 1981; Peters, 1981). In
addition, Ibbotson and Morton (1981) showed that the same was true for
the majority of their left-handed subjects, indicating that this effect was not
simply due to handedness. Byblow et al. (1994) and Byblow and Goodman
(1994) also demonstrated that differences between the hands (left vs. right)
affected performance of the frequency ratios 1: 1 and 1: 2. Based on their
results (obtained for right-handed subjects) they suggested that the right
(preferred) and left (non-preferred) hand can be distinguished with respect
to the degree to which coupling strength scales with movement frequency.
However, in studying the production of polyrhythms, Summers et al.
(1993a) and Summers et al. (1993b) did not observe effects of different
hand arrangements (i.e., either the preferred or the non-preferred hand
performed the faster cadence) on the temporal accuracy of polyrhythmic
240 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

performance. In addition, Peper et al. (in press-b) reported that the

number of transitions to lower-order ratios that were observed during a
multifrequency tapping task did not change when the hand arrangement
was reversed. In line with these results, the present analysis did not show
an effect of hand arrangement on the degree of relaxation in the movement
patterns either. Given our model assumptions this implied that the cou-
pling was not affected by the differences between the left and right hand
per se. The rather consistent lack of influence of hand arrangement on
polyrhythmic (or, more general, multifrequency) performance may indicate
that the effect of the functional roles of the hands (fast vs. slow) on the
asymmetry in coupling is so strong that the differences between the two
hands (left vs. right) are overshadowed.
In general, the peaks in the power spectra were located at integer
multiples of the actual tapping frequency. In a number of bimanual trials,
however, some additional spectral peaks were observed. In the spectra
obtained for the slow hand these additional peaks occurred at odd integer
multiples of 0.5 Xfslow,implying that an increased number of spectral
peaks observed for the slow hand were located at frequencies that coin-
cided with frequencies at which pronounced peaks were observed for the
fast hand (i.e., at integer multiples of f,,,.). Given that the fast hand had
more influence on the slow hand than vice versa, these extra peaks may
have resulted from the coupling between the hands. The relative incidence
of these peaks depended on which hand (left vs. right) performed the faster
cadence. Because this may indicate an effect of hand preference, it seemed
curious that for one of our right-handed drummers (Subject D) the addi-
tional peaks mainly occurred when the left hand performed the faster
frequency, while for subjects A, B, and E the opposite was true (for Subject
C no additional peaks were observed). However, additional tests on hand-
edness, using a shortened Dutch version of the Oldfield Handedness
Questionnaire (Oldfield, 19711, revealed that whereas the other four sub-
jects clearly were right handers (L.Q. ’ ranging from 67% to 86%), Subject
D tended toward ambidexterity (L.Q. = 25%), which might partly account
for the observed reversion of the effect. The extra peaks in the spectra of
the fast hand, as observed for two subjects, were more prominent in the

5
The Laterality Quotient (L.Q.1 is determined by the numbers of Right (R) and Left CL)responses in
the questionnaire: L.Q. = 100% X CR - L)/(R + L) (Oldfield, 19711, resulting in positive values for right
handedness (ranging from 0 to 100%) and negative values for left handedness (ranging from 0 to
- 100%). Larger absolute values reflect stronger lateralization.
 C.E. Peper et al. /Human Movement Science 14 (199s) 217-245 241

slower conditions. This observation may be related to the fact that coupling
strength increased as movement frequency decreased.

4.2. Relative timing and anchoring effects

Analyses of the temporal partitioning of the tapping cycle revealed that

the flight period in the cycle was neither fixed nor a fixed percentage of the
actual overall tapping period. Changes in relaxation can, therefore, not be
attributed to a simple proportional partitioning of the tapping cycle. The
ANOVAs on the relative flight period (i.e., relative to the intertap period)
revealed larger values for the fast hand than for the slow hand. In addition,
the tempo effect was strongest for the fast hand in the bimanual situation.
For the slow hand larger values were observed for unimanual tapping than
for bimanual tapping. The qualitative similarities between these results and
those obtained for the relative contribution of the spectral peak at the
tapping frequency suggest that the relaxation in the oscillations was related
to the period during which the hand dwelled on the tabletop. (Note that a
larger relative flight period is associated with a smaller degree of relax-
ation.)
In the light of our general model, changes in relaxation are considered to
result from the influence of the coupling function which, given the relation
to the observed changes in relative flight period, appears to have a rather
specific phase dependence in the forcing effect. This general observation is
in agreement with the sine circle map in which the coupling function
involves a sinusoidal dependence on the phase of the forced oscillator. The
sine circle map models the behavior of a periodically forced oscillator at
discrete time steps:

where 0 is the phase of the forced oscillator and 0 the ratio between the
uncoupled frequencies. The term (K/2 r)sin 0, represents the forcing
function through which the oscillator is influenced. The forcing influence
depends on the phase of the forced oscillator CO,>. This implies that the
influence of the coupling is not constant throughout the phase evolution of
the forced system, but that it reaches maximal values at the maximal
excursions of the oscillations (at 0 = 0.5~ [mod ~1, corresponding to
x = 0).
Phase dependencies in the effect of forcing have been interpreted by
Beek (1989; see also Beek et al., 1992b) as well as by Kelso and Jeka (1992)
242 C.E. Peper et al. /Human Movement Science I4 (1995) 217-245

as evidence that the essential information for coordination may be confined

to discrete regions in relative phase space. Anchoring effects in the
coordination of rhythmic movements have been reported in a number of
studies (e.g., Beek et al., 1992b; Byblow et al., 1994; Kelso and Jeka, 1992).
In addition Byblow et al. (1994) showed that the most stable performance
in a paced bimanual movement task was obtained when the strongest
anchor point (i.e., maximal pronation as opposed to maximal supination of
the lower arm) was time-locked with the pacing signal. In terms of a
distinction introduced by Bingham (1988) this result was interpreted to
imply that the incidental dynamics (introduced by circumstances associated
with the task) interact with the inherent dynamics (i.e., the dynamics of the
human action system per se). Our results suggest that in tapping the actual
tap (initial contact with the surface) functions as a strong anchor point. The
observed phase dependence in the influence of forcing may be understood
on the basis of two considerations. First, the influence of the nonlinear
coupling is probably largest at the maximal excursions in the movement (cf.
the sine circle map). In addition, one of these two extremes (the actual tap)
is most strongly affected due to the influence of the incidental dynamics
associated with both the pacing signal and the task dynamics (i.e., a tapping
task is defined in terms of the moments in time at which the taps are to be
produced, not in terms of relative phase). These aspects may render this
particular phase in the tapping cycle most susceptible to the coupling
influences.

Acknowledgements

The authors wish to thank Tony van Santvoord and Frank Zaal for their
technical advice and John Whiting and two anonymous reviewers for their
critical comments on an earlier version of this manuscript. This research
was supported by the Netherlands Organization for Scientific Research
(NWO), grant number 575-59-50, and partly supported by NSF Grant SBR
94-22650.

References

Beek, P.J., 1989. Juggling dynamics (Ph.D. thesis). Amsterdam: Free University Press.
Beek, P.J. and W.J. Beek, 1988. Tools for constructing dynamical models of rhythmic movement.
Human Movement Science 7. 301-342.
 C.E. Peper et al. /Human Movement Science 14 (I 995) 217-245 243

Beek, P.J., C.E. Peper and P.C.W. van Wieringen, 1992a. ‘Frequency locking, frequency modulation,
and bifurcations in dynamic movement systems’. In: G.E. Stelmach and J. Requin (Eds.), Tutorials
in motor behavior II (pp. 599-622). Amsterdam: North-Holland.
Beek, P.J., M.T. Turvey and R.C. Schmidt, 1992b. Autonomous and nonautonomous dynamics of
coordinated rhythmic movements. Ecological Psychology 4, 65-95.
Beek, P.J., W.E.I. Rikkert and P.C.W. van Wieringen, in press-a. Limit cycle properties of rhythmic
forearm movements. Journal of Experimental Psychology: Human Perception and Performance.
Beek, P.J., R.C. Schmidt, A.W. Morris, M.-Y. Sim and M.T. Turvey, in press-b. Linear and nonlinear
stiffness and friction in biological rhythmic movements. Biological Cybernetics.
Bingham, G.P., 1988. Task-specific devices and the perceptual bottleneck. Human Movement Science 7.
225-264.
Byblow, W.D., R.G. Carson and D.G. Goodman, 1994. Expressions of asymmetries and anchoring in
bimanual coordination. Human Movement Science 13, 3-28.
Byblow, W.D., and D.G. Goodman, 1994. Performance asymmetries in multifrequency coordination.
Human Movement Science 13, 147-174.
Carson, R.G., W.D. Byblow and D. Goodman, 1993. ‘The dynamical substructure of bimanual
coordination’. In: S.P. Swinnen, H. Heuer, J. Massion and P. Casaer (Eds.), Interlimb coordination:
Neural, dynamical and cognitive constraints (pp. 319-337). Orlando, FL: Academic Press.
Deutsch, D., 1983. The generation of two isochronous sequences in parallel. Perception and Psy-
chophysics 34, 331-337.
Fuchs, A., V.K. Jirsa, H. Haken and J.A.S. Kelso, submitted. Extending the HKB-model of coordinated
movement to oscillators with different eigenfrequencies. Manuscript submitted for publication.
Fuchs, A. and J.A.S. Kelso, 1994. A theoretical note on models of interlimb coordination. Journal of
Experimental Psychology: Human Perception and Performance 20, 1088-1097.
Haken, H., J.A.S. Kelso and H. Bunz, 1985. A theoretical model of phase transitions in human hand
movements. Biological Cybernetics 51, 347-356.
Haken, H., C.E. Peper, P.J. Beek and A. Daffertshofer, in press. A model for phase transitions in
human hand movements during multifrequency tapping. Physica D.
Ibbotson, N.R. and J. Morton, 1981. Rhythm and dominance. Cognition 9, 125-138.
Jagacinski, R.J., E. Marshburn, S.T. Klapp and M.R. Jones, 1988. Test of parallel versus integrated
structure in polyrhythmic tapping. Journal of Motor Behavior 20, 416-442.
Kai, T. and K. Tomita, 1979. Stroboscopic phase portait of a forced nonlinear oscillator. Progress of
Theoretical Physics 61, 54-73.
Kay, B.A., J.A.S. Kelso, E.L. Saltzman and G.S. Schdner, 1987. Space-time behavior of single and
bimanual rhythmical movements: Data and limit cycle model. Journal of Experimental Psychology:
Human Perception and Performance 13, 178-190.
Keith, W.L. and R.H. Rand, 1984. 1: 1 and 1: 2 phase entrainment in a system of two coupled limit
cycle oscillators. Journal of Mathematical Biology 20, 133-152.
Kelso, J.A.S., 1984. Phase transitions and critical behavior in human bimanual coordination. American
Journal of Physiology: Regulatory, Integrative and Comparative Physiology 15, R100&R1004.
Kelso, J.A.S. and G.C. deGuzman, 1988. ‘Order in time: How the cooperation between the hands
informs the design of the brain’. In: H. Haken (Ed.), Neural and synergetic computers (pp.
180-196). Berlin: Springer Verlag.
Kelso, J.A.S., K.G. Holt, P. Rubin and P.N. Kugler, 1981. Patterns of human interlimb coordination
emerge from the properties of non-linear limit cycle oscillatory processes: Theory and data. Journal
of Motor Behavior 13, 226-261.
Kelso, J.A.S. and J.J. Jeka, 1992. Symmetry breaking dynamics of human multilimb coordination.
Journal of Experimental Psychology: Human Perception and Performance 18. 645-668.
244 C.E. Peper et al. /Human Movement Science 14 (1995) 217-245

Kelso, J.A.S. and J.P. Scholz, 1985. ‘Cooperative phenomena in biological motion’. In: H. Haken (Ed.),
Complex systems: Operational approaches in neurobiology, physical systems and computers (pp.
124-149). Berlin: Springer Verlag.
Kelso, J.A.S., J.P. Scholz and G. Schoner, 1986. Nonequilibrium phase transitions in coordinated
biological motion: Critical fluctuations. Physics Letters A 118, 279-284.
Kelso, J.A.S. and G. Schoner, 1988. Self-organization of coordinative movement patterns. Human
Movement Science 7, 27-47.
Klapp, S.T., M.D. Hill, J.G. Tyler, Z.E. Martin, R.J. Jagacinski and M.R. Jones, 1985. On marching to
two different drummers: Perceptual aspects of the difficulties. Journal of Experimental Psychology:
Human Perception and Performance 11, 814-827.
Kopell. N., 1988. ‘Toward a theory of modelling central pattern generators’. In: A.H. Cohen, S.
Rossignol and S. Grillner (Eds.), Natural control of rhythmic movements in vertebrates (pp.
369-413). New York: Wiley.
Kugler, P.N., J.A.S. Kelso and M.T. Turvey, 1980. ‘On the concept of coordinative structures as
dissipative structures: I. Theoretical lines of convergence’. In: G.E. Stelmach and J. Requin (Eds.),
Tutorials in motor behavior (pp. 3-47). Amsterdam: North-Holland.
Miller, N.R., R. Shapiro and T.M. McLaughlin, 1980. A technique for obtaining spatial kinematic
parameters of segments of biomechanical systems from cinematographic data. Journal of Biome-
chanics 13, 535-547.
Oldfield, R.C., 1971. The assessment and analysis of handedness: The Edinburgh inventory. Neuropsy-
chologica 9, 97- 113.
Peper, C.E., P.J. Beek and P.C.W. van Wieringen, 1991. ‘Bifurcations in bimanual tapping: In search of
Farey principles’. In: J. Requin and G.E. Stelmach (Eds.). Tutorials in motor neuroscience (pp.
413-431). Dordrecht: Kluwer.
Peper, C.E., P.J. Beek and P.C.W. van Wieringen, in press-a. Frequency-induced phase transitions in
bimanual tapping. Biological Cybernetics.
Peper, C.E., P.J. Beek and P.C.W. van Wieringen, in press-b. Multifrequency coordination in bimanual
tapping: Asymmetrical coupling and signs of supercriticality. Journal of Experimental Psychology:
Human Perception and Performance.
Peters, M., 1981. Attentional asymmetries during concurrent bimanual performance. Quarterly Journal
of Experimental Psychology 33A, 95-103.
Peters, M. and S. Schwartz, 1989. Coordination of the two hands and effects of attention manipulation
in the production of a bimanual 2:3 polyrhythm. Australian Journal of Psychology 41, 215-224.
Rand, R.H., A.H. Cohen and P.J. Holmes, 1988. ‘Systems of coupled oscillators as models of central
pattern generators’. In: A.H. Cohen, S. Rossignol and S. Grillner (Eds.), Natural control of rhythmic
movements in vertebrates (pp. 333-367). New York: Wiley.
Schmidt, R.C., C. Carello and M.T. Tutvey, 1990. Phase transitions and critical fluctuations in the
visual coordination of rhythmic movements between people. Journal of Experimental Psychology:
Human Perception and Performance 16, 227-247.
Schmidt, R.C., B.K. Shaw and M.T. Turvey, 1993. Coupling dynamics in interlimb coordination. Journal
of Experimental Psychology: Human Perception and Performance 19, 397-415.
Schreiber, I., M. Dolnik, P. Choc and M. Marek. 1988. Resonance behaviour in two-parameter families
of periodically forced oscillators. Physics Letters A 128, 66-70.
Shapiro, R., 1978. Direct linear transformation method for three-dimensional cinematography. The
Research Quarterly 49, 197-205.
Sternad, D., M.T. Turvey and R.C. Schmidt, 1992. Average phase difference theory and 1: 1 entrain-
ment in interlimb coordination. Biological Cybernetics 67, 223-231.
Summers, J.J. and T.M. Kennedy, 1992. Strategies in the production of a 5:3 polyrhythm. Human
Movement Science 11, 101-112.
 C.E. Peper et al. /Human Mouement Science 14 (1995) 217-24.5 245

Summers, J.J., S.K. Ford and J.A. Todd, 1993a. Practice effects on the coordination of the two hands in
a bimanual tapping task. Human Movement Science 12, 111-133.
Summers, J.J., D.A. Rosenbaum, B.D. Burns and S.K. Ford, 1993b. Production of polyrhythms. Journal
of Experimental Psychology: Human Perception and Performance 19, 416-428.
Treffner, P.J. and M.T. Tutvey, 1993. Resonance constraints on rhythmic movement. Journal of
Experimental Psychology: Human Perception and Performance 19, 1221-1237.
Vorberg, D. and R. Hambuch, 1984. Timing of two-handed rhythmic performance. Annals of the New
York Academy of Sciences 423, 390-406.
Wimmers, R.H., P.J. Beek and P.C.W. van Wieringen, 1992. Phase transitions in rhythmic tracking
movements: A case of unilateral coupling. Human Movement Science 11, 217-226.