Delayed auditory feedback in repetitive tapping: A role for the sensory goal

The open-loop model by Wing and Kristofferson has successfully explained many aspects of movement timing. A later adaptation of the model assumes that timing processes do not control the movements themselves, but the sensory consequences of the movements. The present study tested direct predictions from this “sensory-goals model”. In two experiments, participants were instructed to produce regular intervals by tapping alternately with the index fingers of the left and the right hand. Auditory feedback tones from the taps of one hand were delayed. As a consequence, regular intervals between taps resulted in irregular intervals between feedback tones. Participants compensated for this auditory irregularity by changing their movement timing. Compensation effects increased with the magnitude of feedback delay (Experiment 1) and were also observed in a unimanual variant of the task (Experiment 2). The pattern of effects in alternating tapping suggests that compensation processes were anticipatory—that is, compensate for upcoming feedback delay rather than being reactions to delay. All experiments confirmed formal model predictions. Taken together, the findings corroborate the sensory-goals adaptation of the Wing–Kristofferson model.     

Individual differences in timing of discrete and continuous movements: a dimensional approach

This study investigated aspects of individual differences in timing of continuous and discontinuous movements to different pacing signals (auditory or visual), pacing intervals (500, 650, 800, 950 ms), and across effectors (dominant versus non-dominant hand). Correlation and principal component analysis demonstrated that a single statistical dimension accounted for up to 60 % of the explained variance in discontinuous tasks and 25 % of the variance in continuous tasks, when applied to performance obtained from tasks conducted with different effectors and at different pacing rates. Correlation analysis of factor scores representing effector and rate independent task performances showed that timing of discrete or continuous movements can be associated with modality independent mechanisms. Timing variability from discrete and continuous trials was not significantly correlated. This study goes beyond previous correlational work on individual differences in discrete and continuous movements, demonstrating that individual differences in discrete (event-based) or continuous (emergent) motor timing tasks can be modeled as distinctive statistical components with dissimilar capability to capture effector, rate, and modality independent variance.     

Does an auditory distractor sequence affect self-paced tapping?

This study investigated whether an auditory distractor (D) sequence affects the timing of self-paced finger tapping. To begin with, Experiment 1 replicated earlier findings by showing that, when taps are synchronized with an isochronous auditory target (T) sequence, an isochronous D sequence of different tempo and pitch systematically modulates the tap timing. The extent of the modulation depended on the relative intensity of the T and D tones, but not on their pitch distance. Experiment 2 then used a synchronization-continuation paradigm in which D sequences of different tempi were introduced only during continuation tapping. Although the D sequences rarely captured the taps completely, they did increase the tapping variability and deviations from the correct tempo. Furthermore, they eliminated the negative correlation between successive inter-tap intervals and led to intermittent phase locking when the tapping period was close to the period of the D sequence. These distractor effects occurred regardless of whether or not the taps generated auditory feedback tones. The distractor effects thus depend neither on the intention to synchronize with a T sequence nor on the simultaneous perception of two auditory sequences. Rather, they seem to reflect a basic attraction of rhythmic movement to auditory rhythms.     

Coordination dynamics and attentional costs of continuous and discontinuous bimanual circle drawing movements

It has been suggested that the temporal control of rhythmic unimanual movements is different between tasks requiring continuous (e.g., circle drawing) and discontinuous movements (e.g., finger tapping). Specifically, for continuous movements temporal regularities are an emergent property, whereas for tasks that involve discontinuities timing is an explicit part of the action goal. The present experiment further investigated the control of continuous and discontinuous movements by comparing the coordination dynamics and attentional demands of bimanual continuous circle drawing with bimanual intermittent circle drawing. The intermittent task required participants to insert a 400ms pause between each cycle while circling. Using dual-task methodology, 15 right-handed participants performed the two circle drawing tasks, while vocally responding to randomly presented auditory probes. The circle drawing tasks were performed in symmetrical and asymmetrical coordination modes and at movement frequencies of 1Hz and 1.7Hz. Intermittent circle drawing exhibited superior spatial and temporal accuracy and stability than continuous circle drawing supporting the hypothesis that the two tasks have different underlying control processes. In terms of attentional cost, probe RT was significantly slower during the intermittent circle drawing task than the continuous circle drawing task across both coordination modes and movement frequencies. Of interest was the finding that in the intermittent circling task reaction time (RT) to probes presented during the pause between cycles did not differ from the RT to probes occurring during the circling movement. The differences in attentional demands between the intermittent and continuous circle drawing tasks may reflect the operation of explicit event timing and implicit emergent timing processes, respectively.     

Self-induced versus reactive triggering of synchronous movements in a deafferented patient and control subjects

The present study investigates the contribution of tactile-kinesthetic information to the timing of movements. The relative timing of simultaneous tapping movements of finger and foot (hand-foot asynchrony) was examined in a simple reaction time task and in discrete self-initiated taps (Experiment 1), and in externally triggered synchronization tapping (Experiment 2). We compared the performance of a deafferented participant (IW) to the performance of two control groups of different ages. The pattern of results in control groups replicates previous findings: Whereas positive hand-foot asynchronies (hand precedes foot) are observed in a simultaneous reaction to an auditory stimulus, hand-foot asynchronies are negative with discrete self-initiated as well as auditorily paced sequences of synchronized finger and foot taps. In the first case, results are explained by a simultaneous triggering of motor commands. In contrast, self-initiated and auditorily paced movements are assumed to be controlled in terms of their afferent consequences, as provided by tactile-kinesthetic information. The performance of the deafferented participant differed from that of healthy participants in some aspects. As expected on the basis of unaffected motor functions, the participant was able to generate finger and foot movements in reaction to an external signal. In spite of the lack of movement-contingent sensory feedback, the deafferented participant showed comparable timing errors in self-initiated and regularly paced tapping as observed in control participants. However, in discrete self-initiated taps IW's hand-foot asynchronies were considerably larger than in control participants, while performance did not differ from that of controls in continuous movement generation. These findings are discussed in terms of an internal generation of the movement's sensory consequences (forward-modeling).     

Timing variability in circle drawing and tapping: probing the relationship between event and emergent timing

R. Ivry, R. M. Spencer, H. N. Zelaznik, and J. Diedrichsen (2002) have proposed a distinction between timed movements in which a temporal representation is part of the task goal (event timing) and those in which timing properties are emergent. The issue addressed in the present experiment was how timing in conditions conducive to emergent timing becomes established. According to what the authors term the transformation hypothesis, timing initially requires an event-based representation when the temporal goal is defined externally (e.g., by a metronome), but over the first few movement cycles, control processes become established that allow timing to become emergent. Different groups of participants (N = 84) executed either 1 timed interval, 4 timed intervals, or 2 timed intervals separated by a pause. They produced the intervals by either circle drawing, a task associated with emergent timing, or tapping, a task associated with event timing. Analyses of movement variability suggested that similar timing processes were used in the 2 tasks only during the 1st interval. Those results are consistent with the transformation hypothesis and lead to the inference that the transition from event-based control to emergent timing can occur rapidly during continuous movements.     

Timing and the Control of Rhythmic Upper-Limb Movements

Accurate timing of limb displacement is crucial for effective motor control. The authors examined the effects of movement velocity, duration, direction, added mass, and auditory cueing on timing, spatial, and trajectory variability of single- and multijoint rhythmic movements. During single-joint movements, increased velocity decreased timing and spatial variability, whereas increased movement duration increased timing variability but decreased spatial variability. For multijoint movements, regardless of condition, increasing velocity decreased joint timing, spatial, and trajectory variability, but all hand variabilities were unaffected by velocity, duration, load, or direction. Timing, spatial, and trajectory variability was greater at the shoulder compared with the elbow and minimal at the hand, supporting the notion that reaching movements are planned in hand space as opposed to joint space.     

Auditory cueing facilitates temporospatial accuracy of sequential movements

Effectively executing goal-directed behaviours requires both temporal and spatial accuracy. Previous work has shown that providing auditory cues enhances the timing of upper-limb movements. Interestingly, alternate work has shown beneficial effects of multisensory cueing (i.e., combined audiovisual) on temporospatial motor control. As a result, it is not clear whether adding visual to auditory cues can enhance the temporospatial control of sequential upper-limb movements specifically. The present study utilized a sequential pointing task to investigate the effects of auditory, visual, and audiovisual cueing on temporospatial errors. Eighteen participants performed pointing movements to five targets representing short, intermediate, and large movement amplitudes. Five isochronous auditory, visual, or audiovisual priming cues were provided to specify an equal movement duration for all amplitudes prior to movement onset. Movement time errors were then computed as the difference between actual and predicted movement times specified by the sensory cues, yielding delta movement time errors (ΔMTE). It was hypothesized that auditory-based (i.e., auditory and audiovisual) cueing would yield lower movement time errors compared to visual cueing. The results showed that providing auditory relative to visual priming cues alone reduced ΔMTE particularly for intermediate amplitude movements. The results further highlighted the beneficial impact of unimodal auditory cueing for improving visuomotor control in the absence of significant effects for the multisensory audiovisual condition.     

The coupled oscillator model of between-hand coordination in alternate-hand tapping: a reappraisal

Single and alternating hand tapping were compared to test the hypothesis that coordination during rhythmic movements is mediated by the control of specific time intervals. In Experiment 1, an auditory metronome was used to indicate a set of timing patterns in which a 1-s interval was divided into 2 subintervals. Performance, measured in terms of the deviation from the target patterns and variability, was similar under conditions in which the finger taps were made with 1 hand or alternated between the 2 hands. In Experiment 2, the modality of the metronome (auditory or visual) was found to influence the manner in which the produced intervals deviated from the target patterns. These results challenge the notion that bimanual coordination emerges from coupling constraints intrinsic to the 2-hand system. They are in accord with a framework that emphasizes the control of specific time intervals to form a series of well-defined motor events.     

Visuomotor and audiomotor processing in continuous force production of oral and manual effectors

The authors examined force control in oral and manual effectors as a function of sensory feedback (i.e., visual and auditory). Participants produced constant isometric force via index finger flexion and lower lip elevation to 2 force levels (10% and 20% maximal voluntary contraction) and received either online visual or online auditory feedback. Mean, standard deviation, and coefficient of variation of force output were used to quantify the magnitude of force variability. Power spectral measures and approximate entropy of force output were calculated to quantify the structure of force variability. Overall, it was found that the oral effector conditions were more variable (e.g., coefficient of variation) than the manual effector conditions regardless of sensory feedback. No effector differences were found for the structure of force variability with visual or auditory feedback. Oral and manual force control appears to involve different control mechanisms regulating continuous force production in the presence of visual or auditory feedback.     

Mechanisms underlying accuracy in fast goal-directed arm movements in man

This study investigated how accuracy is attained in fast goal-directed arm movements. Subjects were instructed to make arm extension movements over three different distances in random order, with and without visual feedback. Target width was varied proportionally with distance. Movement time was kept as short as possible, but there were well-defined limits with respect to accuracy. There appeared to be a large relative variability (variation coefficient [VC]) in the initial acceleration. The VC in the distance the hand moved during the acceleration phase was much smaller. This reduction was accompanied by a strong negative correlation between the initial acceleration and the duration of the acceleration phase. Further, the VC in the total distance moved was less than the VC in the distance moved during acceleration. This result indicates asymmetry between the acceleration and the deceleration phase. This is confirmed by the negative correlation between the distance the hand moved during acceleration and the distance it moved during deceleration. Withdrawal of visual feedback had a significant effect on movement accuracy. No differences were found in the parameters of the acceleration phase in the two feedback conditions, however. our results point to the existence of a powerful variability compensating mechanism within the acceleration phase. This mechanism seems to be independent of visual feedback; this suggests that efferent information (efference copies) and/or proprioceptive information is/are responsible for the timing of agonist and antagonist activation. The asymmetry between the acceleration and deceleration phase contributes to a reduction in the relative variability in the total distance moved. The fact that the withdrawal of visual feedback affected movement variability only during the deceleration phase indicates that visual information is used in the adjustment of antagonist activity.     

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.     

Auditory information is beneficial for adults with Down syndrome in a continuous bimanual task

Much recent research using discrete unimanual tasks has indicated that individuals with Down syndrome (DS) have more difficulty performing verbal-motor tasks as compared to visual-motor tasks (see Perceptual-Motor Behavior in Down Syndrome, Human Kinetics, Champaign, IL, 2000, p. 305 for a review). In continuous tasks, however, individuals with DS perform better when movement is guided by auditory information compared to visual information (Downs Syndr.: Res. Prac. 4 (1996) 25; J. Sport Exercise Psy. 22 (2000) S90). The aim of the present study was to investigate if there are any differences for adults with DS between visual, auditory and verbal guidance in a continuous bimanual task. Ten adults with DS, 10 adults without DS and 10 typically developing children drew lines bimanually towards the body (down) and away from the body (up) following three different guidance conditions: visual (flashing line), auditory (high tone, low tone), and verbal (“up”, “down”). All participants produced mostly in-phase movements and were close to the 1000 ms target time for all guidance conditions. The adults with DS, however, displayed greater variability in their movement time, movement amplitude and bimanual coordination than adults without DS. For all groups, the left hand was slower and more variable in producing the lateral movements than the right hand. The results regarding guidance information suggest that auditory information is beneficial for repetitive bimanual tasks for adults with DS. Possible mechanisms that cause these results will be discussed.     

On the signals underlying conscious awareness of action

To investigate whether conscious judgments of movement onset are based solely on pre-movement signals (i.e., premotor or efference copy signals) or whether sensory feedback (i.e., reafferent) signals also play a role, participants judged the onset of finger and toe movements that were either active (i.e., self initiated) or passive (i.e., initiated by the experimenter). Conscious judgments were made by reporting the position of a rotating clock hand presented on a computer screen and were then compared to the actual measured time of movement onset. In line with previous studies, judgment errors were found to be anticipatory for both finger and toe movements. There was a significant difference between judgment errors for active and passive movements, with judgments of active movements being more anticipatory than judgments of passive movements. This is consistent with a pre-movement (from here on referred to as an “efferent”) account of action awareness because premotor and efference copy signals are only present in active movements, whereas the main source of movement information in passive movements is sensory feedback which is subject to time delays of conduction (and hence predicts later judgment times for passive movements). However, judgments of active toe movement onset time were less anticipatory than judgments of active finger movement onset time. This pattern of results is not consistent with a pure efferent account of conscious awareness of action onset - as this account predicts more anticipatory judgments for toe movements compared to finger movements. Instead, the data support the idea that conscious judgments of movement onset are based on efferent (i.e., premotor, efference copy) and reafferent (i.e., feedback from the movement) components.     

Periodic change in phase relationship between target and hand motion during visuo-manual tracking task: Behavioral evidence for intermittent control

When one performs visuo-manual tracking tasks, velocity profile of hand movements shows discontinuous patterns even if the target moves smoothly. A crucial factor of this “intermittency” is considerable delay in the sensorimotor feedback loop, and several researchers have suggested that the cause is intermittent correction of motor commands. However, when and how the brain monitors task performance and updates motor commands in a continuous motor task is uncertain. We examined how tracking error was affected by the timing of target disappearance during a tracking task. Results showed that tracking error, defined as the average phase difference between target and hand, varied periodically in all conditions. Hand preceded target at one specific phase but followed it at another, implying that motor control was not performed in a temporally uniform manner. Tracking stability was evaluated by the variance in phase difference, and changed depending on the timing of target-removal. The variability was larger when target disappeared around turning points than that when it disappeared around the center of motion. This shows that visual information at turning points is more effectively exploited for motor control of sinusoidal target tracking, suggesting that our brain controls hand movements with intermittent reference to visual information.     

The Timing Effects of Accent Production in Periodic Finger-Tapping Sequences

When subjects are required to produce short sequences of equally paced finger taps and to accentuate one of the taps, the interval preceding the forceful tap is shortened and the one that immediately follows the accent is lengthened. Assuming that the tapping movements are triggered by an internal clock, one explanation attributes the rnistiming of the taps to central factors: The momentary rate of the clock is accelerated or decelerated as a function of motor preparation to, respectively, increase or decrease the movement force. This hypothesis predicts that the interresponse intervals measured between either tap movement onsets or movement terminations (taps) will show the same timing pattern. A second explanation for the observed interval effects is that the tapping movements are triggered by a regular internal clock but the timing of the successive taps is altered because the forceful movement is completed in less time than the other tap movements are. This "peripheral" hypothesis predicts regular timing of movement onsets but distorted timing of movement terminations. In the present study, the trajectories of the movements performed by subjects were recorded and the interresponse intervals were measured at the beginning and the end of the tapping movements. The results of Experiment 1 showed that neither model can fully explain the interval effects: The fast forceful movements were initiated with an additional delay that took into account the small execution time of these movements. Experiment 2 reproduced this finding and showed that the timing of the onset and contact intervals did not evolve with the repetition of trial blocks. Therefore, the assumption of an internal clock that would trigger the successive movements must be rejected. The results are discussed in the framework of a modified two-stage model in which the internal clock, instead of triggering the tapping movements, provides target time points at which the movements have to produce their meaningful effects, that is, contacts with the response key. The timing distortions are likely to reflect both peripheral and central components.     

The Timing Effects of Accent Production in Periodic Finger-Tapping Sequences

When subjects are required to produce short sequences of equally paced finger taps and to accentuate one of the taps, the interval preceding the forceful tap is shortened and the one that immediately follows the accent is lengthened. Assuming that the tapping movements are triggered by an internal clock, one explanation attributes the mistiming of the taps to central factors: The momentary rate of the clock is accelerated or decelerated as a function of motor preparation to, respectively, increase or decrease the movement force. This hypothesis predicts that the interre-sponse intervals measured between either tap movement onsets or movement terminations (taps) will show the same timing pattern. A second explanation for the observed interval effects is that the tapping movements are triggered by a regular internal clock but the timing of the successive taps is altered because the forceful movement is completed in less time than the other tap movements are. This “peripheral” hypothesis predicts regular timing of movement onsets but distorted timing of movement terminations. In the present study, the trajectories of the movements performed by subjects were recorded and the interresponse intervals were measured at the beginning and the end of the tapping movements. The results of Experiment 1 showed that neither model can fully explain the interval effects: The fast forceful movements were initiated with an additional delay that took into account the small execution time of these movements. Experiment 2 reproduced this finding and showed that the timing of the onset and contact intervals did not evolve with the repetition of trial blocks. Therefore, the assumption of an internal clock that would trigger the successive movements must be rejected. The results are discussed in the framework of a modified two-stage model in which the internal clock, instead of triggering the tapping movements, provides target time points at which the movements have to produce their meaningful effects, that is, contacts with the response key. The timing distortions are likely to reflect both peripheral and central components.     

Goal-directed aiming under restricted viewing conditions with confirmatory sensory feedback

A substantial body of research has examined the speed-accuracy tradeoff captured by Fitts’ law, demonstrating increases in movement time that occur as aiming tasks are made more difficult by decreasing target width and/or increasing the distance between targets. Yet, serial aiming movements guided by internal spatial representations, rather than by visual views of targets have not been examined in this manner, and the value of confirmatory feedback via different sensory modalities within this paradigm is unknown. Here we examined goal-directed serial aiming movements (tapping back and forth between two targets), wherein targets were visually unavailable during the task. However, confirmatory feedback (auditory, haptic, visual, and bimodal combinations of each) was delivered upon each target acquisition, in a counterbalanced, within-subjects design. Each participant performed the aiming task with their pointer finger, represented within an immersive virtual environment as a 1 cm white sphere, while wearing a head-mounted display. Despite visual target occlusion, movement times increased in accordance with Fitts’ law. Though Fitts’ law captured performance for each of the sensory feedback conditions, the slopes differed. The effect of increasing difficulty on movement times was least influential in the haptic condition, suggesting more efficient processing of confirmatory haptic feedback during aiming movements guided by internal spatial representations.     

Aspects of body self-calibration

The representation of body orientation and configuration is dependent on multiple sources of afferent and efferent information about ongoing and intended patterns of movement and posture. Under normal terrestrial conditions, we feel virtually weightless and we do not perceive the actual forces associated with movement and support of our body. It is during exposure to unusual forces and patterns of sensory feedback during locomotion that computations and mechanisms underlying the ongoing calibration of our body dimensions and movements are revealed. This review discusses the normal mechanisms of our position sense and calibration of our kinaesthetic, visual and auditory sensory systems, and then explores the adaptations that take place to transient Coriolis forces generated during passive body rotation. The latter are very rapid adaptations that allow body movements to become accurate again, even in the absence of visual feedback. Muscle spindle activity interpreted in relation to motor commands and internally modeled reafference is an important component in permitting this adaptation. During voluntary rotary movements of the body, the central nervous system automatically compensates for the Coriolis forces generated by limb movements. This allows accurate control to be maintained without our perceiving the forces generated.     

Sensory effects on judgments of short time-intervals

Two experiments were conducted to test the effect of nontemporal factors on duration discrimination. In Exp. 1, a forced-choice adaptive procedure with a standard duration of 400 or 800 ms was employed. It was shown that, for both auditory and visual modes, the discrimination is better with empty intervals (a silent period between two brief signals) than with filled intervals (a continuous signal), but only with shorter durations. In a second experiment, where intervals of the same duration range were employed but were presented with a single-stimulus method, discrimination was better with empty than with filled intervals, and this effect applied to both ranges of duration and both sensory modes. In both experiments, discrimination was better in the auditory than in the visual mode. These data are discussed in the context of current models of timing mechanisms. Received: 16 October 1997 / Accepted: 30 April 1998