Temporal coordination and adaptation to rate change in music performance

People often coordinate their actions with sequences that exhibit temporal variability and unfold at multiple periodicities. We compared oscillator- and timekeeper-based accounts of temporal coordination by examining musicians' coordination of rhythmic musical sequences with a metronome that gradually changed rate at the end of a musical phrase (Experiment 1) or at the beginning of a phrase (Experiment 2). The rhythms contained events that occurred at the same periodic rate as the metronome and at half the period. Rate change consisted of a linear increase or decrease in intervals between metronome onsets. Musicians coordinated their performances better with a metronome that decreased than increased in tempo (as predicted by an oscillator model), at both beginnings and ends of musical phrases. Model performance was tested with an oscillator period or timekeeper interval set to the same period as the metronome (1:1 coordination) or half the metronome period (2:1 coordination). Only the oscillator model was able to predict musicians' coordination at both periods. These findings suggest that coordination is based on internal neural oscillations that entrain to external sequences.     

The acquisition of bimanual coordination is mediated by anisotropic coupling between the hands

The present study was designed to test two predictions from the coupled oscillator model of multifrequency coordination. First, it was predicted that multifrequency tasks that match the inherent manual asymmetry (i.e., the preferred hand assigned to the faster tempo) would be easier to learn than tasks that do not match the inherent dynamics (i.e., the non-preferred hand assigned to the faster tempo). Second, in the latter case acquisition of the multifrequency coordination would involve a reorganisation of the coupling dynamics such that the faster hand would exert a greater influence on the slower hand than vice versa. Sixteen right-handed volunteers received extensive training on a 2:1 coordination pattern involving a bimanual forearm pronation-supination task. Participants were randomly assigned to one of two groups: 1L:2R in which the preferred right hand performed the higher frequency, or 2L:1R in which the non-preferred left hand performed the higher frequency. The dynamic stability of the patterns was assessed by the ability of participants to maintain the coordination pattern as movement frequency was increased. Changes in the directional coupling between the hands was assessed by transition pathways and lead-lag relationship evident in a 1:1 anti-phase frequency-scaled coordination task performed prior to and following three practice sessions of the 2:1 task. The predicted differential stability between the multifrequency patterns was evident in the initial acquisition sessions but by the end of training the two patterns evidenced equivalent stability. Unexpectedly, for both groups the fast hand displayed greater variability in amplitude and movement frequency than the slow hand perhaps reflecting anchoring afforded to the slow hand by synchronising movement endpoints with the auditory pacing metronome. Analysis of pre- to post-training changes to the coupling dynamics in the 1:1 anti-phase task support the hypothesis that acquisition of the 2L:1R pattern involved reorganisation of the inherent dynamics.     

Multilevel coordination stability: integrated goal representations in simultaneous intra-personal and inter-agent coordination

The influence of integrated goal representations on multilevel coordination stability was investigated in a task that required finger tapping in antiphase with metronomic tone sequences (inter-agent coordination) while alternating between the two hands (intra-personal coordination). The maximum rate at which musicians could perform this task was measured when taps did or did not trigger feedback tones. Tones produced by the two hands (very low, low, medium, high, and very high) could be the same as, or different from, one another and the (medium-pitched) metronome tones. The benefits of feedback tones were greatest when they were close in pitch to the metronome and the left hand triggered low tones, while the right hand triggered high tones. Thus, multilevel coordination was facilitated by tones that were easy to integrate with, but perceptually distinct from, the metronome, and by compatibility of movement patterns and feedback pitches.     

Reduced timing variability during bimanual coupling: A role for sensory information

On a repetitive tapping task, the within-hand variability of intertap intervals is reduced when participants tap with two hands as compared to one-hand tapping. Because this bimanual advantage can be attributed to timer variance (Wing-Kristofferson model, 1973a, b), separate timers have been proposed for each hand, whose outputs are then averaged (Helmuth & Ivry, 1996). An alternative notion is that action timing is based on its sensory reafferences (Aschersleben & Prinz, 1995; Prinz, 1990). The bimanual advantage is then due to increased sensory reafference. We studied bimanual tapping with the continuation paradigm. Participants first synchronized their taps with a metronome and then continued without the pacing signal. Experiment 1 replicated the bimanual advantage. Experiment 2 examined the influence of additional sensory reafferences. Results showed a reduction of timer variance for both uni- and bimanual tapping when auditory feedback was added to each tap. Experiment 3 showed that the bimanual advantage decreased when auditory feedback was removed from taps with the left hand. Results indicate that the sensory reafferences of both hands are used and integrated into timing. This is consistent with the assumption that the bimanual advantage is at least partly due to the increase in sensory reafference. A reformulation of the Wing-Kristofferson model is proposed to explain these results, in which the timer provides action goals in terms of sensory reafferences.     

Getting synchronized with the metronome: Comparisons between phase and period correction

Most studies of synchronization have focused on how an established phase relationship between self-produced events (e.g., finger taps) and the clicks of a metronome is maintained when the metronome is regular or subject to unpredictable perturbations. Here we study how synchronization is initially established, using an experimental paradigm in which the metronome is activated after the subject has executed a series of self-paced finger taps. In Exp. 1, the metronome period was constant and equal to the mean of the self-paced inter-response intervals, whereas the initial phase difference of the metronome from the taps varied across trials. The synchronization error patterns could be predicted by a linear phase correction model. Experiment 2 involved both period and phase correction. The initial phase difference was constant, whereas the metronome period varied across trials. The observed synchronization error patterns suggest that the subjects achieved synchronization either by reacting to the second metronome signal or by aiming at the third metronome signal. The pattern of the residual synchronization errors was consistent with the linear phase correction model. These results support the notion that period and phase correction mechanisms are called for by different task variables and contribute differently to sensorimotor synchronization. Received: 20 November 1996 / Accepted: 20 April 1997     

Hierarchical organisation of neuro-anatomical constraints in interlimb coordination

Based on the observation that bimanual finger tapping movements tend toward mirror symmetry with respect to the body midline, despite the synchronous activation of non-homologous muscles, F. Mechsner, D. Kerzel, G. Knoblich, and W. Prinz (2001) [Perceptual basis of bimanual coordination. Nature, 414, 69-73] suggested that the basis of rhythmic coordination is purely spatial/perceptual in nature, and independent of the neuro-anatomical constraints of the motor system. To investigate this issue further, we employed a four finger tapping task similar to that used by F. Mechsner and G. Knoblich (2004) [Do muscle matter in bimanual coordination? Journal of Experimental Psychology: Human Perception and Performance, 30, 490-503] in which six male participants were required to alternately tap combinations of adjacent pairs of index (I), middle (M) and ring (R) fingers of each hand in time with an auditory metronome. The metronome pace increased continuously from 1 Hz to 3 Hz over the course of a 30-s trial. Each participant performed three blocks of trials in which finger combination for each hand (IM or MR) and mode of coordination (mirror or parallel) were presented in random order. Within each block, the right hand was placed in one of three orientations; prone, neutral and supine. The order of blocks was counterbalanced across the six participants. The left hand maintained a prone position throughout the experiment. On the basis of discrete relative phase analyses between synchronised taps, the time at which the initial mode of coordination was lost was determined for each trial. When the right hand was prone, transitions occurred only from parallel symmetry to mirror symmetry, regardless of finger combination. In contrast, when the right hand was supine, transitions occurred only from mirror symmetry to parallel but no transitions were observed in the opposite direction. In the right hand neutral condition, mirror and parallel symmetry are insufficient to describe the modes of coordination since the hands are oriented orthogonally. When defined anatomically, however, the results in each of the three right hand orientations are consistent. That is, synchronisation of finger tapping is determined by a hierarchy of control of individual fingers based on their intrinsic neuro-mechanical properties rather than on the basis of their spatial orientation.     

Temporal stability of rhythmic tapping “on” and “off the beat”: A developmental study

This study, following a dynamic pattern approach, examines age-related differences in the stability of unimanual rhythmic perception-action patterns. Thirty-six children, aged 7, 9, and 11 years, attempted to synchronize their finger tapping to the beats of an auditory metronome, either “on the beat” (i.e., in-phase coordination), or “off the beat” (i.e., antiphase coordination). The temporal stability of these perception- action patterns was measured by the variability of the relative phase between taps and auditory events and by the critical frequency, that is, the frequency at which a loss of stability was observed when the metronome frequency was increased. Age-related differences in stability were found for both relative phase variability and critical frequency. These findings suggest that the relative phase dynamics underlying perception-action coordination patterns change with age in the direction of an increased temporal stability. Received: 29 June 1998 / Accepted: 15 December 1998     

The Dynamics of Bimanual Circle Drawing

A bimanual circle drawing task was employed to elucidate the dynamics of intralimb and interlimb coordination. Right-handed subjects were required to produce circles with both hands in either a symmetrical (mirror) mode (i.e. one hand moving clockwise, the other counter-clockwise) or in an asymmetrical mode (i.e. both hands moving clockwise or counter-clockwise). The frequency of movement was scaled by an auditory metronome from 1.50 Hz to 3.25 Hz in8 (8-sec) steps.In the asymmetrical mode,distortions ofthe movement trajectories, transient departures from the target pattern of coordination, and phase wandering were evident as movement frequency was increased. These features suggested loss of stability. Deviations from circular trajectories were most prominent for movements of the left hand. Transient departures from the required mode of coordination were also largely precipitated by the left hand. The results are discussed with reference to manual asymmetries and mechanisms of interlimb and intersegmental coordination.     

Coordinated flexibility: how initial gaze position modulates eye-hand coordination and reaching

Reaching to targets in space requires the coordination of eye and hand movements. In two experiments, we recorded eye and hand kinematics to examine the role of gaze position at target onset on eye-hand coordination and reaching performance. Experiment 1 showed that with eyes and hand aligned on the same peripheral start location, time lags between eye and hand onsets were small and initiation times were substantially correlated, suggesting simultaneous control and tight eye-hand coupling. With eyes and hand departing from different start locations (gaze aligned with the center of the range of possible target positions), time lags between eye and hand onsets were large and initiation times were largely uncorrelated, suggesting independent control and decoupling of eye and hand movements. Furthermore, initial gaze position strongly mediated manual reaching performance indexed by increments in movement time as a function of target distance. Experiment 2 confirmed the impact of target foveation in modulating the effect of target distance on movement time. Our findings reveal the operation of an overarching, flexible neural control system that tunes the operation and cooperation of saccadic and manual control systems depending on where the eyes look at target onset.     

Phase Transitions and Critical Fluctuations in Rhythmic Coordination of Ipsilateral Hand and Foot

Four subjects performed rhythmic movements of the ankle and the wrist in time with an auditory metronome, in two modes of coordination, antiphase and in-phase. The forearm was placed in either a prone or a supine position. When movements were prepared in the antiphase mode, spontaneous transitions to the in-phase mode, or to phase wandering were observed as metronome frequency was increased. When prepared in the in-phase mode, transitions between in-phase modes or to phase wandering were occasionally observed. Predicted signature features of nonequilbrium phase transitions were noted, including loss of stability and critical fluctuations. The stability of the movement patterns was determined by spatial (dependent upon the direction of movement) rather than anatomical (dependent on the coupling of specific muscle groups) constraints. The position of the forearm had no consistent bearing upon the variability of the phase relations between the limbs, the frequency of phase transitions, or the time of onset of transitions. These results are discussed with reference to the coordination dynamics (e.g., multistability, loss of stability) of multijoint movements.     

Auditory frequency-based inhibition differs from spatial IOR

Uninformative auditory frequency cues have a facilitatory effect on reaction time and accuracy of detection and intensity discrimination of target tones for cue-target intervals of up to 3 sec (Green & McKeown, 2001; Ward, 1997). Under some conditions, however, this facilitatory effect can reverse to an inhibitory effect at cue-target intervals longer than 450 msec (Mondor, Breau, & Milliken, 1998). Thepresent work demonstrates that such inhibitory effects are not found in target-target experiments (Experiment 1) or in cue-target experiments requiring a go-no-go discrimination of the target (Experiment 2), whereas they do appear in the paradigm used by Mondor et al. (1998, Experiment 3), albeit unaffected by the similarity of cue and target. Thus, the frequency-based inhibitory effects sometimes found in auditory cuing tasks can be distinguished empirically from those characterizing spatial inhibition of return (IOR), which are found in both target-target and go-no-go cue-target paradigms. The present work and functional and neurophysiological arguments all support the position that different mechanisms underlie spatial IOR and the inhibitory effects sometimes found in auditory frequency processing.     

Attention as a mediating variable in the dynamics of bimanual coordination

The influence of focal attention on the coordination dynamics in a bimanual circle drawing task was investigated. Six right-handed and seven left-handed subjects performed bimanual circling movements, in two modes of coordination, symmetrical or asymmetrical. The frequency of movement was scaled by an auditory metronome from 1.50 Hz to 3.00 Hz in 7 steps. On each trial, subjects were required to attend either to the dominant hand, to a neutral position, or to the nondominant hand.The uniformity of the relative tangential angle was lower in asymmetrical than in symmetrical conditions, but was not influenced by the direction of attention. In the asymmetrical mode, shifts in RTA relations, suggestive of loss of stability, were evident as the movement frequency was increased. Typically, these shifts were mediated by distortions of the trajectory of the nondominant limb. When the nondominant hand was the focus of attention, movements of this hand were more circular, and temporal variability was reduced, at the cost of a greater deviation from the target frequency. Movements of the dominant hand were not affected by the direction of attention. The findings show that although directed attention acts to modify the coordination dynamics, it does so primarily at the level of the individual hands, rather then in terms of the relation between them.     

To Switch or Not to Switch: Recruitment of Degrees of Freedom Stabilizes Biological Coordination

Recruitment and suppression processes were studied in the swinging-pendulum paradigm (cf. P. N. Kugler & M. T. Turvey, 1987). The authors pursued the hypothesis that active recruitment of previously unmeasured degrees of freedom serves to stabilize an antiphase bimanual coordination pattern and thereby obviates the need for pattern switching from an antiphase to an in-phase coordination pattern, a key prediction of the H. Haken, J. A. S. Kelso, and H. Bunz (1985) model. In Experiment 1, 7 subjects swung single hand-held pendulums in time with an auditory metronome whose frequency increased. Pendulum motion changed from planar (2D) to elliptical (3D), and forearm motion (produced by elbow flexion-extension) was recruited with increasing movement rate for cycling frequencies typically above the pendulum's eigenfrequency. In Experiment 2, 7 subjects swung paired pendulums in either an in-phase or an antiphase coordinative mode as movement rate was increased. With the systematic increase in movement rate, the authors attempted to induce transitions from the antiphase to the in-phase coordinative pattern, with loss of stability the key mechanism of pattern change. Transitions from the antiphase to the in-phase coordinative mode were not observed. Pattern stability, as defined by the variability of the phase relation between the pendulums, was affected only a little by increasing movement rate. As in the single-pendulum case, pendulum motion changed from planar to elliptical, and forearm motion was recruited with increasing cycling frequency. Those results reveal a richer dynamics than previously observed in the pendulum paradigm and support the hypothesis that recruitment processes stabilize coordination in biomechanically redundant systems, thereby reducing the need for pattern switching.     

Effects of visual and auditory guidance on bimanual coordination complexity

Perceptual guidance of movement with simple visual or temporal information can facilitate performance of difficult coordination patterns. Guidance may override coordination constraints that usually limit stability of bimanual coordination to only in-phase and anti-phase. Movement dynamics, however, might not have the same characteristics with and without perceptual guidance. Do visual and auditory guidance produce qualitatively different dynamical organization of movement? An anti-phase wrist flexion and extension coordination task was performed under no specific perceptual guidance, under temporal guidance with a metronome, and under visual guidance with a Lissajous plot. For the time series of amplitudes, periods and relative phases, temporal correlations were measured with Detrended Fluctuation Analysis and complexity levels were measured with multiscale entropy. Temporal correlations of amplitudes and relative phases deviated from the typical 1/f variation towards more random variation under visual guidance. The same was observed for the series of periods under temporal guidance. Complexity levels for all time series were lower in visual guidance, but higher for periods under temporal guidance. Perceptual simplification of the task’s goal may produce enhancement of performance, but it is accompanied by changes in the details of movement organization that may be relevant to explain dependence and poor retention after practice under guidance.     

Evaluation of an imputed pitch velocity model of the auditory tau effect

This article extends an imputed pitch velocity model of the auditory kappa effect proposed by Henry and McAuley (2009a) to the auditory tau effect. Two experiments were conducted using an AXB design in which listeners judged the relative pitch of a middle target tone (X) in ascending and descending three-tone sequences. In Experiment 1, sequences were isochronous, establishing constant fast, medium, and slow velocity conditions. No systematic distortions in perceived target pitch were observed, and thresholds were similar across velocity conditions. Experiment 2 introduced to-be-ignored variations in target timing. Variations in target timing that deviated from constant velocity conditions introduced systematic distortions in perceived target pitch, indicative of a robust auditory tau effect. Consistent with an auditory motion hypothesis, the magnitude of the tau effect was larger at faster velocities. In addition, the tau effect was generally stronger for descending sequences than for ascending sequences. Combined with previous work on the auditory kappa effect, the imputed velocity model and associated auditory motion hypothesis provide a unified quantitative account of both auditory tau and kappa effects. In broader terms, these findings add support to the view that pitch and time relations in auditory patterns are fundamentally interdependent.     

A comparison of visual and auditory short-term memory in monkeys (Cebus apella)

Using a visual and an acoustic sample set that appeared to favour the auditory modality of the monkey subjects, in Experiment 1 retention gradients generated in closely comparable visual and auditory matching (go/no-go) tasks revealed a more durable short-term memory (STM) for the visual modality. In Experiment 2, potentially interfering visual and acoustic stimuli were introduced during the retention intervals of the auditory matching task. Unlike the case of visual STM, delay-interval visual stimulation did not affect auditory STM. On the other hand, delay-interval music decreased auditory STM, confirming that the monkeys maintained an auditory trace during the retention intervals. Surprisingly, monkey vocalizations injected during the retention intervals caused much less interference than music. This finding, which was confirmed by the results of Experiments 3 and 4, may be due to differential processing of “arbitrary” (the acoustic samples) and species-specific (monkey vocalizations) sounds by the subjects. Although less robust than visual STM, auditory STM was nevertheless substantial, even with retention intervals as long as 32 sec.     

Perceptual organization of auditory temporal patterns in European starlings (Sturnus vulgaris)

The perception of continuously repeating auditory patterns by European starlings was explored in seven experiments. In Experiment 1, 4 starlings learned to discriminate between two continuously repeating, eight-element, auditory patterns. Each eight-element pattern was constructed from different temporal organizations of two elements differing in timbre. In Experiments 2–7, the repeating patterns were transformed in ways designed to identify the starlings’ perceptual organization of the patterns. In Experiment 2, the starlings identified patterns beginning with novel starting points. In Experiment 3, discrimination performance was adversely affected by reorganizing the elements in the patterns. In Experiments 4 and 5, the pattern elements were altered. In Experiment 4, the patterns were constructed from two novel elements. In Experiment 5, the temporal location of the two pattern elements was reversed. The transformations of the patterns in Experiments 4 and 5 affected discrimination performance for some, but not all, of the starlings. In Experiments 6 and 7, replacing either of the two elements with silent intervals had no effect on discrimination performance. The results of these experiments identify basic grouping principles that starlings use when they perceive auditory patterns.     

Brief bimanual force pulses: correlations between the hands in force and time.

Three experiments assessed coupling phenomena in the coordination of bimanual force pulses. Experiment 1 required symmetric force pulses (equal target forces and rise times for both hands) using the index finger of each hand. As the authors expected, on the basis of bimanual pointing movement results, this experiment revealed positive correlations between both the force rise times and the force amplitudes of the two hands. Experiments 2 and 3 included asymmetric conditions with different target force amplitudes (Experiment 2) or target rise times (Experiment 3). In Experiment 2 force amplitudes but not rise times were fully decoupled in the asymmetric condition. In the asymmetric condition of Experiment 3, however, neither rise times nor force amplitudes were fully decoupled. The results suggest a hierarchical control structure with temporal control dominating nontemporal control of bimanual force coordination.