Structural Constraints on the Performance of Symmetrical Bimanual Movements with Different Amplitudes

In bimanual movements the amplitude of each hand's movement often depends on the concurrent amplitude of the other hand's movement such that both amplitudes become similar (amplitude coupling). We tested the hypothesis that the strength of amplitude coupling depends on the tempo of performance of a movement sequence, a hypothesis based on a model of bimanual coordination that holds that cross-talk occurs at the execution level as well as at the programming level. Subjects performed bimanual periodic arm movements on two digitizers. In nine conditions constant small, constant large, and alternating small and large amplitudes of each arm were orthogonally combined. Overall tempo was varied by instructing subjects to increase the tempo progressively by 10%. Clear tempo-dependent modulations of the amplitude were observed in movements with instructed constant amplitude when the other hand performed alternating amplitudes. The effect of the size of constant-amplitude movements on the mean amplitude of the other hand indicated cross-talk at the execution level. Cross-talk at the programming level was revealed by the dependence of the current amplitude on the change in the amplitude of the other hand in the preceding cycle. Finally, asymmetric cross-manual effects were observed.     

Static and Phasic Cross-Talk Effects in Discrete Bimanual Reversal Movements

The authors examined the hypothesis that the phasic and the static cross-talk effects found in bimanual movements with different target amplitudes originate at different functional levels of motor control, which implies that the effects can be dissociated experimentally. When the difference between the short and the long amplitudes assigned to the 2 hands of 12 participants was decreased, the static effect disappeared. In contrast, the phasic effect, which can be observed only at short preparation intervals, did not disappear; although it became smaller in absolute terms, in relative terms it did not. In addition, the authors compared the time course of amplitude assimilation with the time course of amplitude variability and examined the correlation between left hand and right hand amplitudes. The disappearance of the phasic amplitude assimilation at increasing preparation intervals turned out to be delayed relative to the decline of the correlation between amplitudes. That finding suggests that the assimilation of mean amplitudes and the correlation between left hand and right hand amplitudes are not fully equivalent indicators of intermanual interactions, but may indicate different kinds of inter-limb coupling.     

Static and phasic cross-talk effects in discrete bimanual reversal movements

The authors examined the hypothesis that the phasic and the static cross-talk effects found in bimanual movements with different target amplitudes originate at different functional levels of motor control, which implies that the effects can be dissociated experimentally. When the difference between the short and the long amplitudes assigned to the 2 hands of 12 participants was decreased, the static effect disappeared, In contrast, the phasic effect, which can be observed only at short preparation intervals, did not disappear; although it became smaller in absolute terms, in relative terms it did not. In addition, the authors compared the time course of amplitude variability and examined the correlation between left hand and right hand amplitudes. The disappearance of the phasic amplitude assimilation at increasing preparation intervals turned out to be delayed relative to the decline of the correlation between amplitudes. That finding suggests that the assimilation of mean amplitudes and the correlation between left hand and right hand amplitudes are not fully equivalent indicators of intermanual interactions, but may indicate different kinds of inter-limb coupling.     

Re-examining structural constraints on the initiation of bimanual movements: the role of starting locations, movement amplitudes, and target locations

Structural constraints affect the coordination of bimanual movements in ways that have been taken to suggest that the specification of different movement amplitudes is subject to strong intermanual interference effects. Most experiments taken to support this notion, however, confounded variations of movement amplitudes with symmetry in starting locations and variations in target location. The present experiment was designed to further investigate the relative influence of the parameters starting location, movement amplitude, and target location on bimanual movement coordination. Participants performed simultaneous reaching movements with the left and right hand from same and different starting locations to same and different target locations. On each trial, two movements could match on none, one, or all of the parameters. We assessed the influence of each parameter by comparing conditions in which only a single parameter matched between the two hands with conditions in which all parameters differed. The reaction-time data revealed some challenging results for previous studies: (1) same starting locations significantly delayed movement initiation; (2) specifying movement amplitudes had virtually no effect on movement initiation, whereas (3) selecting same target locations significantly benefited the bimanual responses. These findings cannot be taken to support the notion that amplitude specification affects the initiation of bimanual movements. Rather, they support the notion that the initial starting locations of the two hands and the selection of target locations decide about the ease with which we perform bimanual reaching movements.     

The control of amplitude and direction in a bimanual coordination task

Bimanual coordination requires task-specific control of the spatial and temporal characteristics of the movements of both hands. The present study focused on the spatial relationship between hand movements when their amplitude and direction were manipulated. In the experiment in question, participants were instructed to draw two lines simultaneously. These two lines were instructed to be drawn in mirror symmetric or perpendicular directions of each other while the length was instructed to be the same or different. The coordinative quality of amplitude control was compared when the task required symmetric and asymmetric bimanual spatial coordination patterns. Results showed that the amplitude accuracy decreased when different amplitudes and/or directions had to be generated simultaneously. The coordinative quality of direction was also compared when the task required symmetric and asymmetric bimanual spatial coordination patterns. Unlike amplitude, the direction accuracy was largely independent of coordination symmetry/asymmetry of direction or amplitude. The results suggest that the coordinative quality of amplitude control does not only interfere with amplitude asymmetry, but it also interferes with direction asymmetry. Moreover, in bimanual coordination amplitude control is more vulnerable to the influence of direction control demands than vice versa.     

Intermanual cross-talk effects in unimanual choice reactions

Intermanual interactions originate at different levels of motor control. Interactions during specification of movement characteristics should affect reaction time for choice between left-hand and right-hand movements. In two experiments combinations of short and long target amplitudes for reversal movements of the left and right hand were cued with variable precueing intervals. Upon presentation of the response signal a unimanual left-hand or right-hand movement had to be produced. Reaction time was faster when same target amplitudes were precued than when different target amplitudes were. At short precueing intervals the longer reaction time with different target amplitudes (early effect) was accompanied by an amplitude assimilation: Short amplitudes were too long, and long amplitudes were too short. At longer precueing intervals the longer reaction time with different target amplitudes (late effect) was accompanied by a higher choice accuracy. These findings are taken to indicate a transient parametric coupling of amplitude specifications, which produces the early and the late effects by way of different mechanisms-namely different degrees of advance specification and generalized de-coupling, which affects the process of choice between hands.     

Specification of movement amplitudes for the left and right hands: evidence for transient parametric coupling from overlapping-task performance

Bimanual coordination tasks suggest transient cross-talk between concurrent specification processes for movements of the left and right hand that vanishes as the time for specification increases. In 2 experiments with overlapping and successive unimanual tasks, the hypothesis of transient coupling was examined for a psychological-refractory-period paradigm. Time for specification was manipulated by varying the delay between first and second signal (Experiment 1) and by precuing the first response (Experiment 2). Participants performed rapid reversal movements of same or different amplitudes with the left and right hands. With different amplitudes, reaction times (RTs) of the second responses were longer than with same amplitudes at short delays, and this disappeared at longer delays in Experiment 1. In Experiment 2, precuing also reduced the difference between RTs of second responses in same-amplitude and different-amplitude trials. These findings are consistent with the hypothesis of transient coupling during amplitude specification obtained with bimanual tasks.     

Intermanual Interactions During Simultaneous Execution and Programming of Finger Movements

In bimanual movements, some differences between the movements performed by the two hands cause interference, while others do not. Similarly, in choice between responses with the left and right hand some differences between the two movements increase RT, while others do not. It is suggested that both kinds of effects are, at least in part, due to the incompatibility between processes that determine characteristics of movements jointly for both hands and that are present during preparation as well as during execution. This hypothesis implies that during execution of one movement, programming of a different movement to be performed with the other hand should be impaired, as compared to a condition in which the successive movements of both hands are the same. This expectation was confirmed for finger movements of different forms where an effect on choice RT had been shown previously. On the other hand, interference between execution and programming is not to be expected when successive movements differ in characteristics that are likely to be specified separately for each hand, as indicated by a lacking effect in choice experiments. This expectation was confirmed for successive movements performed with different fingers of either hand as compared to movements performed with the same fingers.     

Intermanual interactions during simultaneous execution and programming of finger movements

In bimanual movements, some differences between the movements performed by the two hands cause interference, while others do not. Similarly, in choice between responses with the left and right hand some differences between the two movements increase RT, while others do not. It is suggested that both kinds of effects are, at least in part, due to the incompatibility between processes that determine characteristics of movements jointly for both hands and that are present during preparation as well as during execution. This hypothesis implies that during execution of one movement, programming of a different movement to be performed with the other hand should be impaired, as compared to a condition in which the successive movements of both hands are the same. this expectation was confirmed for finger movements of different forms where an effect on choice RT had been shown previously. On the other hand, interference between execution and programming is not to be expected when successive movements differ in characteristics that are likely to be specified separately for each hand, as indicated by a lacking effect in choice experiments. This expectation was confirmed for successive movements performed with different fingers of either hand as compared to movements performed with the same fingers.     

Bimanual interference associated with the selection of target locations

Four experiments were conducted to identify the locus of interference observed during the preparation of bimanual reaching movements. Target locations were specified by color, and the right-hand and left-hand targets could be either the same or a different color. Movements of different amplitudes (Experiment 1) or different directions (Experiment 2) to targets of the same color were initiated more quickly than symmetric movements to targets of different colors. These results indicate that costs observed during bimanual movements arise during target selection rather than during motor programming. Experiments 3 and 4 further examined the interference associated with target selection. Reaction time costs were found with unimanual movements when the target was presented among distractors associated with responses for the other hand. Interference observed during bimanual reaching appears to reflect difficulty in segregating the response rules assigned to each hand.     

Intermanual interactions related to movement amplitudes and endpoint locations

In almost all studies of bimanual movements with same and different amplitudes, the difference between amplitudes has been confounded with a difference between endpoint locations. The present authors varied those parameters orthogonally. In addition, they presented target locations on the surface on which the movements were produced (direct cues) and on a monitor (indirect cues). Participants' (N = 12) reaction times were longer when both amplitudes and endpoint locations differed than when they were the same. Intermanual amplitude correlations were reduced whenever 1 of the movement parameters differed for the 2 hands; only when cues were presented on the monitor was the amplitude correlation further reduced when both movement parameters were different. The results indicate that structural constraints on bimanual movements take effect on both amplitudes and endpoint locations. The relative importance of those 2 parameters is largely independent of the type of cue.     

Hand preference, practice order, and spatial assimilations in rapid bimanual movement

When subjects make rapid bimanual aiming movements over different distances, spatial assimilations are shown; the shorter distance limb overshoots when paired with a longer distance limb. Recent research has also shown spatial assimilations to be greater in the nonpreferred left limb of right-handed subjects, but it is not known whether the increased spatial assimilations represent a handedness effect or one of hemispheric lateralization of motor control. To determine the nature of the asymmetric effect, left- (n = 32) and right- (n = 60) handed subjects part practiced, then whole practiced, short (20 degrees ) and long 60 degrees ) reversal movements. During whole practice, both groups showed spatial assimilations in the shorter distance limb, particularly when the left limb performed the short movement. This asymmetry was greatest for right-handed subjects, but left-handed subjects showed smaller, but systematic effects, providing moderate support for the hypothesis that the asymmetric effect is due to hemispheric lateralization of motor control. All interlimb differences in spatial accuracy for the short and long movements were eliminated with practice, however, suggesting the asymmetric effect was temporary as well. In addition, subjects who part practiced the long movement just prior to whole practice showed greater overshooting in the short distance limb compared with subjects who followed the other practice order throughout whole practice and the no-KR retention trials. Such findings suggest that the part-practice order of bimanual tasks can directionally bias whole-task performance.     

The influence of movement cues on intermanual interactions

In two experiments, we studied intermanual interactions in bimanual reversal movements and bimanual aiming movements. Targets were presented on a monitor or directly on the table on which the movements were produced. Amplitudes for each hand were cued symbolically or spatially either in advance of an imperative signal or simultaneous with it. In contrast to findings of Diedrichsen et al. (Psychological Science, 12, 493–498, 2001), reaction times for different-amplitude movements were longer than for same-amplitude movements both for symbolic and spatial cues presented on the monitor and directly on the table. However, with symbolic cues the effect of the relation between target amplitudes was considerably stronger than with spatial cues, no matter where the cues were presented. Intermanual correlations of amplitudes, movement times, and reaction times were smaller with different than with same target amplitudes, and this modulation was more pronounced when targets and cues were presented on the monitor than when they were presented on the table. The findings are taken to suggest that the basic reaction-time disadvantage of different-amplitude movements results from interference between concurrent processes of amplitude specification. Additional factors like interference between concurrent processes of mapping cues on movement characteristics may add strongly to it.     

Manual asymmetries in bimanual reaching: the influence of spatial compatibility and visuospatial attention

The goal of the present investigation was to explore the possible expression of hemispheric-specific processing during the planning and execution of a bimanual reaching task. Participants (N = 9) completed 80 bimanual reaching movements (requiring simultaneous, bilateral production of arm movements) to peripherally presented targets while selectively attending to either their left or right hand. Further, targets were presented in spatially compatible (ipsilateral to the aiming limb) and incompatible (contralateral to the aiming limb) response contexts. It was found that the left hand exhibited temporal superiority over the right hand in the response planning phase of bimanual reaching, indicating a left hand/right hemisphere advantage in the preparation of a bimanual response. During response execution, and consistent with the view that interhemispheric processing time (Barthelemy & Boulinguez, 2002) or biomechanical constraints (Carey, Hargreaves, & Goodale, 1996) generate temporal delays, longer movement times were observed in response to spatially incompatible target positions. However, no hemisphere-specific benefit was demonstrated for response execution. Based on these findings, we propose lateralized processing is present at the time of response planning (i.e., left hand/right hemisphere processing advantage); however, lateralized specialization appears to be annulled during dynamic execution of a bimanual reaching task.     

Cognitive constraints on motor imagery

Executed bimanual movements are prepared slower when moving to symbolically different than when moving to symbolically same targets and when targets are mapped to target locations in a left/right fashion than when they are mapped in an inner/outer fashion [Weigelt et al. (Psychol Res 71:238–447, 2007)]. We investigated whether these cognitive bimanual coordination constraints are observable in motor imagery. Participants performed fast bimanual reaching movements from start to target buttons. Symbolic target similarity and mapping were manipulated. Participants performed four action conditions: one execution and three imagination conditions. In the latter they indicated starting, ending, or starting and ending of the movement. We measured movement preparation (RT), movement execution (MT) and the combined duration of movement preparation and execution (RTMT). In all action conditions RTs and MTs were longer in movements towards different targets than in movements towards same targets. Further, RTMTs were longer when targets were mapped to target locations in a left/right fashion than when they were mapped in an inner/outer fashion, again in all action conditions. RTMTs in imagination and execution were similar, apart from the imagination condition in which participants indicated the start and the end of the movement. Here MTs, but not RTs, were longer than in the execution condition. In conclusion, cognitive coordination constraints are present in the motor imagery of fast (<1600 ms) bimanual movements. Further, alternations between inhibition and execution may prolong the duration of motor imagery.     

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.     

Bimanual coupling effects during arm immobilization and passive movements

When humans simultaneously perform different movements with both hands, each limb movement interferes with the contralateral limb movement (bimanual coupling). Previous studies on both healthy volunteers and patients with central or peripheral nervous lesions suggested that such motor constraints are tightly linked to intentional motor programs, rather than to movement execution. Here, we aim to investigate this phenomenon, by using a circles-lines task in which, when subjects simultaneously draw lines with the right hand and circles with the left hand, both the trajectories tend to become ovals (bimanual coupling effect). In a first group, we immobilized the subjects’ left arm with a cast and asked them to try to perform the bimanual task. In a second group, we passively moved the subjects’ left arm and asked them to perform voluntary movements with their right arm only. If the bimanual coupling arises from motor intention and planning rather than spatial movements, we would expect different results in the two groups. In the Blocked group, where motor intentionality was required but movements in space were prevented by immobilization of the arm, a significant coupling effect (i.e., a significant increase of the ovalization index for the right hand lines) was found. On the contrary, in the Passive group, where movements in space were present but motor intentionality was not required, no significant coupling effect was observed. Our results confirmed, in healthy subjects, the central role of the intentional and predictive operations, already evidenced in pathological conditions, for the occurrence of bimanual coupling.     

Intermovement interval and spatial assimilation effects in sequential bimanual and unimanual movements

This study extended earlier work by showing spatial assimilations in sequential bimanual and unimanual movements separated by 1.5-3.5 s. In Experiments 1 and 2, 30 right-handed participants (18-22 years of age) made rapid single and bimanual lever reversals of 20 degrees and 60 degrees assigned to 1.5, 2.5, or 3.5 s intermovement interval groups. Participants self-timed the intermovement interval in the first experiment, but it was provided in the second experiment using separate auditory stimuli. In the third experiment, participants performed both the 20 degree and 60 degree movement with the same hand. In all experiments, the shorter-distance limb overshot and the longer-distance limb undershot the targets in both bimanual and unimanual sequential movements relative to single movements in all three intermovement interval groups, particularly in the non-dominant left limb. The results suggest that assimilation effects in sequential movements are caused by command interactions at the planning level, but the effects are reduced by practice.     

Separate movement planning and spatial assimilation effects in sequential bimanual aiming movements

This study extended earlier work by showing spatial assimilations in sequential bimanual aiming movements when the participant preplanned only the first movement of a two-movement sequence. Right-handed participants (n=20, aged 18 to 22 years) made rapid lever reversals of 20 degrees and 60 degrees singly and sequentially with an intermovement interval of 2.5 sec. Following blocked single practice of both movements in each hand (15 trials each), two sets of 30 sequential practice trials were completed. The sequences began with either the long or the short movement and the participant always knew the goal of the first movement. During the intermovement interval, the experimenter gave instructions to complete the sequence with a short movement, a long movement, or no movement in a random order. Compared to the single trials, both movements in the sequence overshot the short-distance and undershot the long-distance goal. Spatial errors increased when a change in the movement goal was required for the second movement in the sequence. The experiment demonstrated that separate planning of sequential aiming movements can reduce spatial assimilation effects, but interference due to practice organization and switching the task's goal must also be overcome in order to produce accurate aiming movements.     

Asynchrony in discrete bimanual aiming: Evidence for visual strategies of coordination

The bimanual coupling literature supposes an inherent drive for synchrony between the upper limbs when making discrete bimanual movements. The level of synchrony is argued to be task dependent, reliant on the visual demands of the two targets, and the result of a complex pattern of hand and eye movements (Bingham, Hughes, & Mon-Williams, 2008 ; Riek, Tresilian, Mon-Williams, Coppard, & Carson, 2003 ). However, recent work by Bruyn and Mason ( 2009 ) suggests that temporal coordination is not solely influenced by visual saccades. In this experimental series, a total of 8 participants performed congruent movements to targets either near or far from the midline. Targets far from the midline, requiring a visual saccade, resulted in greater terminal asynchrony. Initial and terminal asynchrony were not consistent, but linked to the task demands at that stage of the movement. If the asynchrony evident at the end of a bimanual movement is due to a complex pattern of hand and eye movements then the removal of visual feedback should result in an increase in synchrony. Sixteen participants then completed congruent and incongruent bimanual aiming movements to near and/or far targets. Movements were made with or without visual feedback of hands and targets. Analyses revealed that movements made without visual feedback showed increased synchrony between the limbs, yet movements to incongruent targets still showed greater asynchrony. We suggest that visual constraints are not the sole cause of asynchrony in discrete bimanual movements.