Catching With Both Hands: An Evaluation of Neural Cross-Talk and Coordinative Structure Models of Bimanual Coordination

Kelso, Southard, and Goodman (1979) and Marteniuk and MacKenzie (1980) have each proposed a different theoretical model for bimanual coordination. In the model of Kelso et al., a close temporal relationship between the hands in a bimanual task is predicted, even when each hand is required to move different distances. In Marteniuk and MacKenzie's model, separate motor commands are issued so that each limb will arrive simultaneously at the specified movement endpoint, leading to low temporal associations between limbs. In most previous work on bimanual coordination, manual aiming tasks with differing constraints have been used by subjects in individual studies. In this study, the usefulness of existing models for predicting performance in a real-world catching task in which the required movement pattern was constrained by ball flight characteristics was examined. E1even university students caught tennis balls with both hands in the following 3 conditions: Condition 1. Ball projected to the right shoulder area (left hand moved a greater distance than the right); Condition 2. Ball projected to center of the chest area, (both hands moved same distance); and Condition 3. Ball projected to left shoulder area (right hand moved a greater distance). Kinematic data (time to peak velocity, movement initiation time) indicating significant cross-correlations between the left and right limbs in all 3 conditions concurred with the data of Kelso et al. (1979) on manual aiming. Timing appeared to be an essential variable coordinating bimanual interceptive actions. Although the limbs moved at different speeds when each was required to move different distances, times to peak velocity showed strong associations, suggesting the presence of a coordinative structure.     

Bimanual movement control: Information processing and interaction effects

Three experiments were designed to investigate the underlying processes in bimanual control. With one hand alone, or with both simultaneously, subjects moved styli from the midline of the body to lateral targets as quickly and accurately as possible. The distance moved and the weight of the styli were varied. Results of reaction time, movement time, and kinematic trajectory analyses question the conclusions of Kelso, Southard and Goodman (1979) regarding the synchronicity of movement of the two limbs. Temporal parameters for the two limbs indicated marked departures from synchronicity, and there was evidence for a left-right asymmetry. The dependent variables of movement time and constant error indicated that there was interaction between the two limbs. The results are discussed in terms of three postulated processes underlying bimanual movement: limb selection (one or two), specification of movement locations and the specification of movement intensities.     

Spatial Conceptual Influences on the Coordination of Bimanual Actions: When a Dual Task Becomes a Single Task

When the left and right hands produce 2 different rhythms simultaneously, coordination of the hands is difficult unless the rhythms can be integrated into a unified temporal pattern. In the present study, the authors investigated whether a similar account can be applied to the spatial domain. Participants (N = 8) produced a movement trajectory of semicircular form in single-limb and bimanual conditions. In the bimanual tasks, 1 limb moved above the other in the frontal plane. Bimanual unified tasks were constructed so that the spatial paths to be produced by the 2 limbs could be easily conceptualized as parts of a unified circle pattern. Bimanual distinct tasks availed a less obvious spatial pattern that would unify the 2 tasks. Performance of the spatial patterns was more accurate in the unified task, despite similar demands placed on the coordination dynamics between the limbs in the 2 cases (e.g., the phase relations). The authors conclude that a dual task becomes a single task, and interlimb interference is reduced, when the spatial patterns produced by the 2 hands form a geometric arrangement that can be conceptualized as a unified representation.     

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.     

Timing and co-ordination of repetitive bimanual movements

Similar timing of movements of the two hands has been observed when they are moved to separate targets (Kelso et al., 1979). This was taken as evidence for a low-level, co-ordinative structure that constrains the muscles of the arms to function as a single unit.

An experiment to investigate the relation between voluntary timing control and timing in bimanual movement is described. The task required subjects to make repetitive movements of unequal difficulty for the two hands with the hands arriving synchronously at their respective targets. Estimates of the covariance of successive intervals defined by pairs of left-right responses (arrivals at the targets) were not negative. It is shown that this indicates that the motor delay between the timer regulating repetition rate and the overt responses has no component common to left- and right-responses. Although the co-ordinative structure is described as low-level, in terms of the time sequence of operations associated with each response pair, the data indicate its place is before, not after, the timer.     

Roles of Visual Information for Control of Reaching Movements in Children

Poststroke hemiparetic individuals (n = 9) and a control group (n = 9) completed a frequency-scaled circle-drawing task in unimanual and bimanual conditions. Measures of intralimb spatial and temporal task accuracy and interlimb coordination parameters were analyzed. Significant reductions in task performance were seen in both limbs of the patients and controls with the introduction of bimanual movement. Spatial performance parameters suggested that the 2 groups focused on different hands during bimanual conditions. In the controls, interlimb coordination variables indicated predictable hand dominance effects, whereas in the patient group, dominance was influenced by the side of impairment and prior handedness of the individual. Therefore, in this particular bimanual task, performance improvements in the hemiplegic side could not be elicited. Intrinsic coupling asymmetries between the hands can be altered by unilateral motor deficits.     

Spatial conceptual influences on the coordination of bimanual actions: when a dual task becomes a single task

When the left and right hands produce 2 different rhythms simultaneously, coordination of the hands is difficult unless the rhythms can be integrated into a unified temporal pattern. In the present study, the authors investigated whether a similar account can be applied to the spatial domain. Participants (N = 8) produced a movement trajectory of semicircular form in single-limb and bimanual conditions. In the bimanual tasks, 1 limb moved above the other in the frontal plane. Bimanual unified tasks were constructed so that the spatial paths to be produced by the 2 limbs could be easily conceptualized as parts of a unified circle pattern. Bimanual distinct tasks availed a less obvious spatial pattern that would unify the 2 tasks, despite similar demands placed on the coordination dynamics in the 2 cases (e.g., the phase relations). The authors conclude that a dual task becomes a single task, and interlimb interference is reduced, when the spatial patterns produced by the 2 hands form a geometric arrangement that can be conceptualized as a unified representation.     

Co-ordination of bimanual movements in a centrally deafferented patient executing open loop reach-to-grasp movements

Many everyday tasks require that we use our hands co-operatively. For tasks where both hands are required to perform the same action, a common motor program can be used. But, where each hand must perform a different action, some degree of independent control of each hand is required. In this paper we examine the co-ordination of bimanual movement kinematics in a female patient recovering from brain injury involving anterior regions of the parietal lobe of the right hemisphere, which has resulted in a dense hemianaesthesia of her left arm. A particular focus of this paper is the co-ordination of bimanual movements for reaches executed without visual feedback. Specifically we present new data, which quantify the synchronisation of patient D.B.'s hands by comparing their relative time lag at the start and the end of her bimanual reaches. The results are discussed with particular reference to the role played by limb proprioception in the planning and control of prehension movements.     

Central fatigue mechanisms are responsible for decreases in hand proprioceptive acuity following shoulder muscle fatigue

Muscle fatigue is a complex phenomenon, consisting of central and peripheral mechanisms which contribute to local and systemic changes in motor performance. In particular, it has been demonstrated that afferent processing in the fatigued muscle (e.g., shoulder), as well as in surrounding or distal muscles (e.g., hand) can be altered by fatigue. Currently, it is unclear how proximal muscle fatigue affects proprioceptive acuity of the distal limb. The purpose of the present study was to assess the effects of shoulder muscle fatigue on participants’ ability to judge the location of their hand using only proprioceptive cues. Participants’ (N = 16) limbs were moved outwards by a robot manipulandum and they were instructed to estimate the position of their hand relative to one of four visual reference targets (two near, two far). This estimation task was completed before and after a repetitive pointing task was performed to fatigue the shoulder muscles. To assess central versus peripheral effects of fatigue on the distal limb, the right shoulder was fatigued and proprioceptive acuity of the left and right hands were tested. Results showed that there was a significant decrease in the accuracy of proprioceptive estimates for both hands after the right shoulder was fatigued, with no change in the precision of proprioceptive estimates. A control experiment (N = 8), in which participants completed the proprioceptive estimation task before and after a period of quiet sitting, ruled out the possibility that the bilateral changes in proprioceptive accuracy were due to a practice effect. Together, these results indicate that shoulder muscle fatigue decreases proprioceptive acuity in both hands, suggesting that central fatigue mechanisms are primarily responsible for changes in afferent feedback processing of the distal upper limb.     

When two limbs are weaker than one: Sensorimotor syncopation with alternating hands

This study addresses the demands of alternating bimanual syncopation, a coordination mode in which the two hands move in alternation while tapping in antiphase with a metronomic tone sequence. Musically trained participants were required to engage in alternating bimanual syncopation and five other coordination modes: unimanual syncopation where taps are made (with the left or right hand) after every tone; unimanual syncopation where taps are made after every other tone; bimanual synchronization with alternating hands; unimanual synchronized tapping with every tone; and unimanual tapping with every other tone. Variability in tap timing was greatest overall for alternating bimanual syncopation, indicating that it is the most difficult. This appears to be due to instability arising from the simultaneous presence of two levels of antiphase coordination (one between the pacing sequence and the hands, the other between the two hands) rather than factors relating to movement frequency or dexterity limits of the nonpreferred hand.     

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.     

Bimanual reaching across the hemispace: Which hand is yoked to which?

When both hands perform concurrent goal-directed reaches, they become yoked to one another. To investigate the direction of this coupling (i.e., which hand is yoked to which), the temporal dynamics of bimanual reaches were compared with equivalent-amplitude unimanual reaches. These reaches were to target pairs located on either the left or right sides of space; meaning that in the bimanual condition, one hand's contralateral (more difficult) reach accompanied by the other hand's ipsilateral (easier) reach. By comparing which hand's difficult reach was improved more by the presence of the other hand's easier ipsilateral reach, we were able to demonstrate asymmetries in the coupling. When the cost of bimanual reaching was controlled for the contralateral reaching left hand's performance was improved, suggesting that the left hand is yoked to the right during motor output. In contrast, the right hand showed the greatest improvements for contralateral reaching in terms of reaction time, pointing toward a dominant role for the left hand in the processes prior to movement onset. The results may point toward a mechanism for integrating the unitary system of attention with bimanual coordination.     

Subcortical locus of temporal coupling in the bimanual movements of a callosotomy patient

The timing of repetitive movements was assessed in a callosotomy patient under unimanual and bimanual conditions. Similar to neurologically healthy individuals, the patient exhibited strong temporal coupling in the bimanual condition. Moreover, for both the left and right hands, within-hand temporal variability was reduced in the bimanual condition compared to the unimanual conditions. This bimanual advantage is hypothesized to reflect the temporal integration of separable timing signals, one associated with the left hand and one associated with the right hand (Helmuth, L. L., & Ivry, R. B. (1996). When two hands are better than one: Reduced timing variability during bimanual movements. Journal of Experimental Psychology: Human Perception and Performance, 2, 278–293). The fact that it persists following callosotomy is inconsistent with models that attribute bimanual coordination in these patients to the control of a single hemisphere. Rather, the results suggest that motor commands from the two hemispheres are integrated subcortically.PsychINFO Classification: 2330; 2340; 2520     

Bilateral associations of low-level unilateral performance: an unremarked aspect of limb control

The experimenters examine upper limb movement discrimination performance in an arm-raising task for bilateral associations of low-level unilateral performance. On a cue from the experimenter, young adults (n = 23) with no history of shoulder injury raised either their left arm, right arm, or both together in a forward flexion movement until their hand or hands contacted an unseen, adjustable, overhead stop. The participant then judged which of the 5 possible stop positions, in the 12-20 degrees range forward of true vertical, the participant had contacted on the particular trial. Results showed that for the 16 participants whose best performance was in 1 of the unimanual conditions discrimination scores in the bilateral condition were equivalent to those of their worse-performing limb. For the 7 participants whose best discrimination performance was obtained on bimanual arm-raising, scores for the 2 unimanual conditions were equivalently low. Therefore, when a single limb that can perform well operates in conjunction with a limb performing at a lower level, the consequence is lowering the bimanual movement discrimination performance.     

Functional coupling between the limbs during bimanual reach-to-grasp movements

While it is frequently advantageous to be able to use our hands independently, many actions demand that we use our hands co-operatively. In this paper we present two experiments that examine functional binding between the limbs during the execution of bimanual reach-to-grasp movements. The first experiment examines the effect of gaze direction on unimanual and bimanual reaches. Even when subjects' eye movements are restricted during bimanual reaches so that they may only foveate one target object, the limbs remain tightly synchronized to a common movement duration. In contrast, grip aperture is independently scaled to the size of the target for each hand. The second experiment demonstrates however, that the independent scaling of grip aperture is task dependent. If the two target objects are unified so that they appear to be part of a single object, grip apertures become more similar across the hands (i.e., grip aperture to the large target object is reduced in size while peak aperture to the small target item is increased in size). These results suggest that the coupling of the limbs can operate at a functional level.     

Bimanual and Unimanual Convergent Goal-Directed Movement Times

Three experiments are reported, investigating the effects of using 1 or 2 hands when making convergent low index of difficulty (ID) and visually controlled movements (2 hands meeting together). The experiments involved movements in four different cases—a probe held in the right hand and moved to a target held in the stationary left hand, vice versa of this arrangement, both hands moving with the probe in the right hand and target in the left hand, and vice-versa of this arrangement. Experiments were the standard Fitts’ paradigm, moving a pin into a hole and a low-ID task. In Fitts’ task, 2-hand movements were faster than 1 hand only at higher IDs; this was also the case in the pin-to-hole transfer task and the movement times were lower when the pin was held in the preferred hand. Movements made with low ID showed a small effect of 1- or 2-handed movements, with the effective amplitude of the movement being reduced by about 20% when 2 hands were used.     

Spatial topological constraints in a bimanual task.

Previous research has shown that the concurrent performance of two manual tasks results in a tight temporal coupling of the limbs. The intent of the present experiment was to investigate whether a similar coupling exists in the spatial domain. Subjects produced continuous drawing of circles and lines, one task at a time or bimanually, for a 20 s trial. In bimanual conditions in which subjects produced the circle task with one hand and the line task with the other, there was a clear tendency for the movement path of the circle task to become more line-like and the movement path of the line task to become more circle-like, i.e., a spatial magnet effect. A bimanual circle task and a bimanual line task did not exhibit changes in the movement path when compared to single-hand controls. In all bimanual conditions, the hands were tightly temporally locked. The evidence of temporal coupling and concomitant accommodation in the movement path for the conditions in which the hands were producing different shapes suggests that spatial constraints play a role in the governance of bimanual coordinated actions.     

Asymmetric control of force and symmetric control of timing in bimanual finger tapping

An experiment was conducted to examine the control of force and timing in bimanual finger tapping. Participants were trained to produce both unimanual (left or right hand) and bimanual finger-tapping sequences with a peak force of 200 g and an intertap interval (ITI) of 400 ms. During practice, visual force feedback was provided pertaining to the hand performing the unimanual tapping sequences and to either the dominant or the nondominant hand in the bimanual tapping sequences. After practice, the participants produced the learned unimanual and bimanual tapping sequences in the absence of feedback. In those trials the force produced by the dominant (right) hand was significantly larger than that produced by the nondominant (left) hand, in the absence of a significant difference between the ITIs produced by both hands. Furthermore, after unilateral feedback had been provided of the force produced by the nondominant hand, the force output of the dominant hand was significantly more variable than that of the nondominant hand. In contrast, after feedback had been provided of the force produced by the dominant hand, the variability of the force outputs of the two hands did not differ significantly. These results were discussed in the light of both neurophysiological and anatomical findings, and were interpreted to imply that the control of timing (in bimanual tasks) may be more tightly coupled in the motor system than the control of force.     

Dynamical aspects of learning an interlimb rhythmic movement pattern

Learning a bimanual rhythmic task is explored from the perspective that motor skill acquisition involves the successive reparameterization of a dynamical control structure in the direction of increasing stability, where the intentional process of reparameterization is itself dynamical. Subjects learned to oscillate pendulums held in the right and left hands such that the right hand frequency was twice that of the left (2:1 frequency lock). Over 12 learning sessions of 20 trials each, we interpreted the decreasing fluctuations in the frequency locking to be an index of the increasing concavity of the underlying potential, a measure of stability; the time required to achieve the 2: 1 pattern was interpreted as indexing the relaxation time of an intentional dynamic. Power spectral analyses of the phase velocity ratio exhibited two strategies for acquiring the interlimb movement pattern: (a) adding spectral peaks at integer multiples of the left hand frequency or (b) distributing power across many frequencies in a l/f-like manner. Results are discussed in terms of the promise of a dynamical approach to learning coordinated movements.     

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.