Andras Semjen

Callosotomy patients exhibit temporal uncoupling during continuous bimanual movements

Rhythmic bimanual movements are highly constrained in the temporal domain, with the gestures of the two hands tightly synchronized. Previous studies have implicated a subcortical locus for temporal coupling based on the observation that these constraints persist in callosotomy patients. We now report that such coupling is restricted to movements entailing a discrete event (such as a movement onset). Three callosotomy patients exhibited a striking lack of temporal coupling during continuous movements, with the two hands oscillating at non-identical frequencies. We propose a subcortical locus of temporal coupling for movements involving discrete events. In contrast, synchronization between the hands during continuous movements depends on interhemispheric transmission across the corpus callosum.

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.

Simulating a neural cross-talk model for between-hand interference during bimanual circle drawing.

Abstract. Studies on drawing circles with both hands in the horizontal plane have shown that this task is easy to perform across a wide range of movement frequencies under the symmetrical mode of coordination, whereas under the asymmetrical mode (both limbs moving clockwise or counterclockwise) increases in movement frequency have a disruptive effect on trajectory control and hand coordination. To account for these interference effects, we propose a simplified computer model for bimanual circle drawing based on the assumptions that (1) circular trajectories are generated from two orthogonal oscillations coupled with a phase delay, (2) the trajectories are organized on two levels, “intention” and “motor execution”, and (3) the motor systems controlling each hand are prone to neural cross-talk. The neural cross-talk consists in dispatching some fraction of any force command sent to one limb as a mirror image to the other limb. Assuming predominating coupling influences from the dominant to the nondominant limb, the simulations successfully reproduced the main characteristics of performance during asymmetrical bimanual circle drawing with increasing movement frequencies, including disruption of the circular form drawn with the nondominant hand, increasing dephasing of the hand movements, increasing variability of the phase difference, and occasional reversals of the movement direction in the nondominant limb. The implications of these results for current theories of bimanual coordination are discussed.

Planning and Timing of Finger-Tapping Sequences with a Stressed Element

Advance planning and execution-time organization of sequences of five finger taps were studied in four experiments. Intertap intervals were required to be equal. In some experimental conditions, one of the taps had to be stronger than the other four. Serial position of the stressed tap, number of alternative stress positions, and tapping rate were manipulated. Time to initiate the sequence after presentation of a reaction stimulus (RT), intertap intervals, and force of the taps were measured. the different effects of stress production and choosing between alternative stress locations on the RT of fast as compared to slow sequences suggest that a plan was selected and activated for the whole sequence only when it had to be executed at a fast rate. Additional organization of the fast sequences during execution was inferred from the intertap intervals, force patterns, and stress location errors, that were all different from those observed in slow sequences. The effects of stress production on timing are discussed in relation to existing timing models.

A Cognitive Neuroscience Perspective on Bimanual Coordination and Interference

We argue that bimanual coordination and interference depends critically on how these actions are represented on a cognitive level. We first review the literature on spatial interactions, focusing on the difference between movements directed at visual targets and movements cued symbolically. Interactions manifest during response planning are limited to the latter condition. These results suggest that interactions in the formation of the trajectories of the two hands are associated with processes involved in response selection, rather than interactions in the motor system. Neuropsychological studies involving callosotomy patients argue that these interactions arise from transcallosal interactions between cortically-based spatial codes. The second half of the chapter examines temporal constraints observed in bimanual movements. We propose that most bimanual movements are marked by a common event structure, an explicit representation that ensures temporal coordination of the movements. The translation of an abstract event structure into a movement with a particular timing pattern is associated with cerebellar function, although the resulting temporal coupling during bimanual movements may be due to the operation of other subcortical mechanisms. For rhythmic movements that do not entail an event structure, timing may be an emergent property. Under such conditions, both spatial and temporal coupling can be absent. The emphasis on abstract levels of constraint makes clear that limitations in bimanual coordination overlap to a considerable degree with those observed in other domains of cognition.

Attention as a mediating variable in the dynamics of bimanual coordination

Abstract 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.

Distributed planning of movement sequences.

Three experiments are reported that examine whether fast finger-tapping sequences are entirely planned before execution starts (advance planning), or if they can be started while planning is still under way (distributed planning). Subjects performed finger tapping sequences of three to eight taps at a high rate, under both simple and 2-choice reaction time (RT) conditions. The sequences differed in the location of an accentuated element within them. The RT to choose between sequences with different accent locations progressively decreased as an inverse function of the time-distance between the initial tap and the first point at which the alternative sequences differed. The shortening in choice reaction time (CRT) was never accompanied by noticeable changes in the inter-response times or force patterns of the tapping sequences. The RT to initiate sequences with accent location known beforehand (SRT condition) showed, in two of three experiments, a weak decreasing trend as the accentuated tap shifted away from the beginning of the sequence. The SRT results suggest a possible predominance of advance planning when the same sequence is repeated over a series of trials. The CRT results are taken as evidence that planning of the sequence beyond the unpredictable tap could be distributed before and after sequence initiation. Several factors are discussed that may influence the balance between planning in advance of, and planning in parallel with, sequence execution.

On controlling force and time in rhythmic movement sequences: the effect of stress location.

Accentuation involves modulation of motor intensity. It differentiates a movement from others within a motor sequence. Does the serial position of the accent characterize the whole sequence as a particular response? How are the control of time and force coordinated in the motor sequence? Subjects produced sequences of four fingertaps on a key. Time of onset and force of each tap were recorded. Tapping rate was imposed by a string of four clicks delivered at 180-msec intervals before each trial. A flashed digit served as go signal. It indicated to the subject which of the four taps had to be tapped stronger (stress +) or weaker (stress -) than all the others. These conditions were run in separate series. Reaction time (RT) of the sequence increased when the number of equally likely locations of the stress increased from 2 to 4. RT was also longer under the stress - than under the stress + condition. Tapping intervals were longer before and after the stressed tap than elsewhere in the series. The first and last intervals tended to be longer than the second one. These effects were the same under both stress conditions. The RT data indicate that the motor sequence is identified as a particular response before it starts. Timing is partly force-independent, but is modulated by central processes that control force.

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 models 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.

On the Timing Basis of Bimanual Coordination in Discrete and Continuous Tasks

Motor events are behaviorally meaningful, discrete entities (e.g., key strokes) that are generated at some specific portion of an effectors movement trajectory. Bimanual coordination may be conceptualized with reference to such discrete motor events or with reference to continuous movement trajectories. Studies inspired by the former approach suggest that hand coordination is primarily achieved by assigning a coherent timing goal structure to the motor events produced by each hand. Studies conducted with the latter approach have shown that between-hand interdependence may also arise from the cross-coupling of the command signals that generate each hands motion. Little is known, however, about the relationships between timing-level coordination and trajectory-level coordination of the hands. Some aspects of these relationships are analyzed using data from experiments that involved bimanual finger tapping and circle drawing at identical and different frequencies.