II Sensory feedback influences on keytapping motor tasks

Subjects were trained to tap a key continuously at a specific rate, and with a specific amount of pressure (regularity task). Performance of this task was studied under conditions of: (a) decreased auditory feedback (masking noise through earphones), (b) decreased visual feedback (tapping hand screened from view), (c) vibration (vibrators applied to forearm in order to “mask” proprioceptive feedback), (d) digital block of tapping finger, and (e) combination of all four conditions. Significant changes in rate and intensity of tapping resulted under conditions of decreased auditory feedback, vibration, and the combined condition.

In the second part of the study, the effects of different delayed sensory events on keytapping were examined. The five conditions of delayed sensory feedback were: (a) delayed auditory feedback, (b) delayed visual feedback, (c) delayed tactile feedback, (d) the first three delayed sensory events presented simultaneously, and (e) condition (d) repeated with digital block of the tapping finger.

The conditions of delayed sensory feedback did not markedly alter performance of the regularity task. The same conditions of delayed sensory feedback did, however, produce highly significant changes in the performance of a more complex pattern task. All of these delay conditions produced parallel changes in the pattern task, namely increased intensity and decreased rate of tapping. The fact that the pattern task is more disturbed by delayed sensory feedback than the regularity task suggests that temporal complexity of the task is one determinant of the degree to which it will be disturbed by a delay in sensory feedback.     

Having a body versus moving your body: How agency structures body-ownership

We investigated how motor agency in the voluntary control of body movement influences body awareness. In the Rubber Hand Illusion (RHI), synchronous tactile stimulation of a rubber hand and the participant's hand leads to a feeling of the rubber hand being incorporated in the participant's own body. One quantifiable behavioural correlate of the illusion is an induced shift in the perceived location of the participant's hand towards the rubber hand. Previous studies showed that the induced changes in body awareness are local and fragmented: the proprioceptive drift is largely restricted to the stimulated finger. In the present study, we investigated whether active and passive movements, rather than tactile stimulation, would lead to similarly fragmented body awareness. Participants watched a projected image of their hand under three conditions: active finger movement, passive finger movement, and tactile stimulation. Visual feedback was either synchronous or asynchronous with respect to stimulation of the hand. A significant overall RHI, defined as greater drifts following synchronous than asynchronous stimulation, was found in all cases. However, the distribution of the RHI across stimulated and non-stimulated fingers depended on the kind of stimulation. Localised proprioceptive drifts, specific to the stimulated finger, were found for tactile and passive stimulation. Conversely, during active movement of a single digit, the proprioceptive drifts were not localised to that digit, but were spread across the whole hand. Whereas a purely proprioceptive sense of body-ownership is local and fragmented, the motor sense of agency integrates distinct body-parts into a coherent, unified awareness of the body.     

Synchronizing actions with events: The role of sensory information

Tasks requiring the subject to tap in synchrony to a regular sequence of stimulus events (e.g., clicks) usually elicit a response pattern in which the tap precedes the click by about 30-50 msec. This “negative asynchrony” was examined, first, by instructing subjects to use different effectors for tapping (hand vs. foot; Experiments 1 and 2), and second, by administering extrinsic auditory feedback in addition to the intrinsic tactile/kinesthetic feedback (Experiment 2). Experiment 3 controlled whether the results observed in Experiment 2 were due to purely sensory factors within the auditory modality. Results suggest that taps are synchronized with clicks at the central level by superimposing two sensory codes in time: the tactile/kinesthetic code that represents the tap (the afferent movement code) and the auditory code that represents the click (the afferent code that results from the guiding signal). Because the processing times involved in code generation are different for these two central codes, the tap has to lead over the click.     

A note on time-frequency analysis of finger tapping

Finger tapping involves 3 important features: time, spatial amplitude, and frequency. In classical analysis, investigators examine timing parameters; in spectral analysis, they examine frequency parameters. Both types of analysis are based on stationary tap information. The authors propose that time-frequency analysis is a useful tool for analyzing nonstationary finger tapping. They describe the method and give examples of frequency modulation, age difference, and speed transition that demonstrate additional insights one can gain by using this analysis.     

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.     

Synchronization in repetitive smooth movement requires perceptible events

Accurate timing performance during auditory–motor synchronization has been well documented for finger tapping tasks. It is believed that information pertaining to an event in movement production aids in detecting and correcting for errors between movement cycle completion and the metronome tone. Tasks with minimal event-related information exhibit more variable synchronization and less rapid error correction. Recent work from our laboratory has indicated that a task purportedly lacking an event structure (circle drawing) did not exhibit accurate synchronization or error correction (Studenka & Zelaznik, in press). In the present paper we report on two experiments examining synchronization in tapping and circle drawing tasks. In Experiment 1, error correction processes of an event-timed tapping timing task and an emergently timed circle drawing timing task were examined. Rapid and complete error correction was seen for the tapping, but not for the circle drawing task. In Experiment 2, a perceptual event was added to delineate a cycle in circle drawing, and the perceptual event of table contact was removed from the tapping task. The inclusion of an event produced a marked improvement in synchronization error correction for circle drawing, and the removal of tactile feedback (taking away an event) slightly reduced the error correction response of tapping. Furthermore, the task kinematics of circle drawing remained smooth providing evidence that event structure can be kinematic or perceptual in nature. Thus, synchronization and error correction, characteristic of event timing (Ivry, Spencer, Zelaznik, & Diedrichsen, 2002; Repp, 2005), depends upon the presence of a distinguishable source of sensory information at the timing goal.     

The contribution of tactile reafference to temporal regularity during bimanual finger tapping

In a repetitive tapping task, the within-hand variability of intertap intervals is reduced when participants tap with both hands, as opposed to single-handed tapping. This bimanual advantage can be attributed to timer variance (according to the Wing-Kristofferson model). Separate timers have been proposed for each hand whose outputs are then averaged (Helmuth & Ivry, 1996, Journal of Experimental Psychology: Human Perception and Performance, 22, 278-293). Alternatively, timing might be based on sensory reafference and the bimanual advantage due to the enhancement of sensory reafferences. This alternative hypothesis was tested in three experiments. In the first experiment, we replicated the bimanual advantage in tapping with two fingers of the same hand compared with single finger tapping. In the second experiment, we demonstrated that the bimanual advantage decreased when tactile reafferences from left-hand taps were omitted (by contact-free tapping). In the third experiment, participants tapped bimanually with the index fingers of both hands firmly mechanically coupled. The bimanual advantage was replicated for this condition. Results are consistent with the assumption that the bimanual advantage is due to the sensory reafferences of the second hand. We suggest that our results are best explained by a reformulation of the Wing-Kristofferson model, in which the timer provides action goals in terms of sensory reafferences.     

Crossing the arms confuses the clocks: Sensory feedback and the bimanual advantage

The bimanual advantage refers to the finding that tapping with two fingers on opposite hands exhibits reduced timing variability, as compared with tapping with only one finger. Two leading theories propose that the bimanual advantage results from the addition of either sensory (i.e., enhanced feedback) or cognitive (i.e., multiple timekeeper) processes involved in timing. Given that crossing the arms impairs perception of tactile stimuli and modulates cortical activation following tactile stimulation, we investigated the role of crossing the arms in the bimanual advantage. Participants tapped unimanually or bimanually with their arms crossed or uncrossed on a tabletop or in the air. With arms crossed, we expected increased interval timing variance. Similarly, for air tapping, we expected reduced bimanual advantage, due to reduced sensory feedback. A significant bimanual advantage was observed for the uncrossed, but not the crossed posture in tabletop tapping. Furthermore, removing tactile feedback from taps eliminated the bimanual advantage for both postures. Together, these findings suggest that crossing the arms likely impairs integration of internal (i.e., effector-specific) and external (i.e., environment-specific) information and that this multisensory integration is crucial to reducing timing variability during repetitive coordinated bimanual tasks.     

Auditory-motor coupling of bilateral finger tapping in children with and without DCD compared to adults

The ability to modulate bilateral finger tapping in time to different frequencies of an auditory beat was studied. Twenty children, 7 years of age, 10 with and 10 without developmental coordination disorder (DCD), and 10 adults tapped their left index and right middle fingers in an alternating pattern in time with an auditory signal for 15s (four trials each, randomly, at 0.8, 1.6, 2.4, 3.2 Hz per finger). Dominant and non-dominant finger data were collapsed since no differences emerged. All three groups were able to modulate their finger frequency across trials to closely approximate the signal frequency but children with DCD were unable to slow down to the lowest frequency. Children with DCD were more variable in tap accuracy (SD of relative phase) and between finger coordination than typically developing children who were respectively more variable than the adults. Children with DCD were unable to consistently synchronize their finger with the beat. Adults were tightly synchronized and often ahead of the beat while children without DCD tended to be behind the beat. Overall, these results indicated that children with DCD can only broadly match their finger movements to an auditory signal with variability and poor synchronicity as key features of their auditory-fine-motor control. Individual inspection of the data revealed that five children with DCD had difficulty matching the slowest frequencies and that these children also had higher variability and lower percentile MABC scores from the movement assessment battery for children (MABC) than other children with DCD. Three children with DCD were more variable only at higher frequencies and two performed like typically developing children.     

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.     

Vision,proprioception and corollary discharge in a movement recall test

Movement recall was investigated in relation to the sensory processes involved in a triangle drawing task.Forty subjects in two groups, one with and one without visual feedback, performed a recall task involving movements of their index finger. All subjects attended different experimental sessions in which (1) all proprioceptive feedback was eliminated by the ischaemic block technique, (2) muscle spindle feedback was distorted by vibration of the muscles and tendons involved in the movement, and (3) proprioceptive feedback was normal.Within each session subjects were required firstly to recall triangular movements made for them passively by the experimenter, and secondly, to recall movements they had made actively. Results indicated comparable accuracy in recall of active movements in all conditions, and a decrement in passive recall dependent on the availability of the alternative sources of feedback. The results indicated a process of integrated contribution of all inputs to the perception of movement; redundancy in information when all channels are available; and a role of corollary discharge in recall of movements.     

Common pathways in mental imagery and pain perception: an fMRI study of a subject with an amputated arm

The present paper reviews data from two previous studies in our laboratory, as well as some additional new data, on the neuronal representation of movement and pain imagery in a subject with an amputated right arm. The subject imagined painful and non-painful finger movements in the amputated stump while being in a MRI scanner, acquiring EPI-images for fMRI analysis. In Study I (Ersland et al., 1996) the Subject alternated tapping with his intact left hand fingers and imagining "tapping" with the fingers of his amputated right arm. The results showed increased neuronal activation in the right motor cortex (precentral gyrus) when tapping with the fingers of the left hand, and a corresponding activation in the left motor cortex when imagining tapping with the fingers of the amputated right arm. Finger tappings of the intact left hand fingers also resulted in a larger activated precentral area than imagery "finger tapping" of the amputated right arm fingers. In Study II (Rosen et al., 2001 in press) the same subject imagining painful and pleasurable finger movements, and still positions of the fingers of the amputated arm. The results showed larger activations over the motor cortex for movement imagining versus imagining the hand being in a still position, and larger activations over the sensory cortex when imagining painful experiences. It can therefore be concluded that not only does imagery activate the same motor areas as real finger movements, but also that adding instructions of pain together with imaging moving the fingers intensified the activation compared with adding instructions about non-painful experiences. From these studies, it is clear that areas activated during actual motor execution to a large extent also are activated during mental imagery of the same motor commands. In this respect the present studies add to studies of visual imagery that have shown a similar correspondence in activation between actual object perception and imagery of the same object.     

Timing goals in bimanual coordination

Phase coupling between movement trajectories has been proposed as the basic mechanism of hand coordination in the production of bimanual rhythmic movements with a 1:2 frequency ratio. Here a central temporal coupling view is proposed as an alternative. Extending previous models of two-handed synchronic and alternate-hand tapping, we hypothesized that 1:2 tapping is performed under the control of a single internal timekeeper set at the frequency required for the fast hand. The fast hand is assumed to use every signal and the slow hand every other signal of the timekeeper, to produce actions coordinated in time. The model's predictions for the variance-covariance pattern of tap timing within and across hands were tested in an experiment that required tapping with both hands with 1:1 or 1:2 frequency ratio. The finger contact on the response plate was to be short or long, according to instruction. Prolonged finger contact entailed profound modifications in the movement trajectories but failed to modify the variance-covariance pattern of the tap timing. This pattern proved to conform to predictions under both the short and the long contact conditions, thus supporting the central temporal coupling hypothesis.     

Reduced corticomotor excitability with cyclic passive movement: a study using transcranial magnetic stimulation

Human voluntary movement involves the integration of kinaesthetic information with efferent motor activity during the planning and execution stages of movement. While much is known of the inhibitory and excitatory effects resulting from activation of specific kinaesthetic sensory receptors, in the present study we employed cyclic passive movement of the index finger in order to activate a range of kinaesthetic receptors in a manner that was intended to correspond to how these receptors might be active during a comparable voluntary movement. We intended to identify how this passive movement protocol might affect the excitability of the corticomotor pathway. During 1 Hz cyclic passive movement of the index finger there was an approximately 60% reduction in the amplitude of the motor evoked response from the first dorsal interosseous muscle. The results of the present study demonstrate that passive movement can have a profound effect on the excitability of the corticomotor pathway.     

The influence of speed and force on bimanual finger tapping patterns.

The current study investigated factors that affect the stability of anti-phase bimanual finger tapping. Past research employing the order parameter and control parameter concepts, has identified frequency of movement as a control parameter that affects the stability of finger movement patterns (the order parameter). The present study investigated the hypothesis that multiple movement related variables can interact to influence the stability of an order parameter. Specifically, the combined effect of the rate of movement and movement force on the stability of bimanual finger tapping was examined. Participants were required to initiate an anti-phase tapping pattern under three different movement rate conditions (600, 400, and 200 ms), and were required to increase the force of one finger at the onset of a randomly presented stimulus. The results indicate that an increase in the force parameter at lower tapping rates (600 ms) did not affect the phase relation of the fingers, however at higher rates (200 and 400 ms), the introduction of a force parameter resulted in fluctuations of the phase relation of the fingers, which were followed by pattern shifts from anti-phase to in-phase tapping. The results indicate that movement force and rate of movement interact to influence the outcome of the tapping pattern. Further research is required to investigate force as a control parameter.     

Cooperative selection of movements: The optimal selection model

How one selects a movement when faced with alternative ways of doing a task is a central problem in human motor control. Moving the fingertip a short distance can be achieved with any of an infinite number of combinations of knuckle, wrist, elbow, shoulder, and hip movements. The question therefore arises: how is a unique combination chosen? In our model, choice is achieved by consideration of the similarity between the task requirements and the optimal biomechanical performance of each limb segment. Two variants of the model account for the movements that are selected when subjects freely oscillate the fingertip and when they tap against an obstacle. An important feature of both is that the impulse of collision with an obstacle (as in drumming with the hand or tapping with the finger) is assumed to be controlled in part by aiming for a point beyond the surface being struck. Thus, a force-related control variable may be represented and controlled spatially.     

Body movement enhances the extraction of temporal structures in auditory sequences

Auditory and motor systems interact in processing auditory rhythms. This study investigated the effect of intuitive body movement, such as head nodding or foot tapping, on listeners’ ability to entrain to the pulse of an auditory sequence. A pulse-finding task was employed using an isochronous sequence of tones in which tones were omitted at pseudorandom positions. Musicians and non-musicians identified their subjectively fitting pulse either using periodic body movement or through listening only. The identified pulse was measured subsequently by finger tapping. Movement appeared to assist pulse extraction especially for non-musicians. The chosen pulse tempi tended to be faster with movement. Additionally, movement led to higher synchronization stabilities of the produced pulse along the sequence, regardless of musical training. These findings demonstrated the facilitatory role of body movement in entraining to auditory rhythms and its interaction with musical training.     

Rhythms and responses

Rhythms are fundamental to behavior, but the control mechanism for timed responses is not known. Many theorists have assumed that there is a central clock coordinating behavior in all sensory modalities and response modes. We tested this hypothesis using a rhythmic tapping task in which university undergraduates first attempted to synchronize responses with brief auditory, tactile, or visual stimuli and then continued to tap at the same rate on their own. Performance was most variable with visual stimuli and least variable with auditory stimuli. The detailed results suggest that performances are not based on a common clock, but, rather, different strategies are employed when the task is presented in different modalities. We reject the hypothesis of a single timing mechanism as controlling behavior and, in doing so, question the validity of information processing models that are formulated without regard to temporal relations among their conjectured processes.     

The role of emotional context in facilitating imitative actions

In two experiments, we explored whether emotional context influences imitative action tendencies. To this end, we examined how emotional pictures, presented as primes, affect imitative tendencies using a compatibility paradigm. In Experiment 1, when seen index finger movements (lifting or tapping) and pre-instructed finger movements (tapping) were the same (tapping–tapping, compatible trials), participants were faster than when they were different (lifting–tapping, incompatible trials). This compatibility effect was enhanced when the seen finger movement was preceded by negative primes compared with positive or neutral primes. In Experiment 2, using only negative and neutral primes, the influence of negative primes on the compatibility effect was replicated with participants performing two types of pre-instructed finger movements (tapping and lifting). This emotional modulation of the compatibility effect was independent of the participants' trait anxiety level. Moreover, the emotional modulation pertained primarily to the compatible conditions, suggesting facilitated imitation due to negatively valent primes rather than increased interference. We speculate that negative stimuli increase imitative tendencies as a natural response in potential flight-or-fight situations.     

Spatial constraints on visual-tactile cross-modal distractor congruency effects

Across three experiments, participants made speeded elevation discrimination responses to vibrotactile targets presented to the thumb (held in a lower position) or the index finger (upper position) of either hand, while simultaneously trying to ignore visual distractors presented independently from either the same or a different elevation. Performance on the vibrotactile elevation discrimination task was slower and less accurate when the visual distractor was incongruent with the elevation of the vibrotactile target (e.g., a lower light during the presentation of an upper vibrotactile target to the index finger) than when they were congruent, showing that people cannot completely ignore vision when selectively attending to vibrotactile information. We investigated the attentional, temporal, and spatial modulation of these cross-modal congruency effects by manipulating the direction of endogenous tactile spatial attention, the stimulus onset asynchrony between target and distractor, and the spatial separation between the vibrotactile target, any visual distractors, and the participant’s two hands within and across hemifields. Our results provide new insights into the spatiotemporal modulation of crossmodal congruency effects and highlight the utility of this paradigm for investigating the contributions of visual, tactile, and proprioceptive inputs to the multisensory representation of peripersonal space.