Hierarchical organisation of neuro-anatomical constraints in interlimb coordination

Based on the observation that bimanual finger tapping movements tend toward mirror symmetry with respect to the body midline, despite the synchronous activation of non-homologous muscles, F. Mechsner, D. Kerzel, G. Knoblich, and W. Prinz (2001) [Perceptual basis of bimanual coordination. Nature, 414, 69-73] suggested that the basis of rhythmic coordination is purely spatial/perceptual in nature, and independent of the neuro-anatomical constraints of the motor system. To investigate this issue further, we employed a four finger tapping task similar to that used by F. Mechsner and G. Knoblich (2004) [Do muscle matter in bimanual coordination? Journal of Experimental Psychology: Human Perception and Performance, 30, 490-503] in which six male participants were required to alternately tap combinations of adjacent pairs of index (I), middle (M) and ring (R) fingers of each hand in time with an auditory metronome. The metronome pace increased continuously from 1 Hz to 3 Hz over the course of a 30-s trial. Each participant performed three blocks of trials in which finger combination for each hand (IM or MR) and mode of coordination (mirror or parallel) were presented in random order. Within each block, the right hand was placed in one of three orientations; prone, neutral and supine. The order of blocks was counterbalanced across the six participants. The left hand maintained a prone position throughout the experiment. On the basis of discrete relative phase analyses between synchronised taps, the time at which the initial mode of coordination was lost was determined for each trial. When the right hand was prone, transitions occurred only from parallel symmetry to mirror symmetry, regardless of finger combination. In contrast, when the right hand was supine, transitions occurred only from mirror symmetry to parallel but no transitions were observed in the opposite direction. In the right hand neutral condition, mirror and parallel symmetry are insufficient to describe the modes of coordination since the hands are oriented orthogonally. When defined anatomically, however, the results in each of the three right hand orientations are consistent. That is, synchronisation of finger tapping is determined by a hierarchy of control of individual fingers based on their intrinsic neuro-mechanical properties rather than on the basis of their spatial orientation.     

Does handedness determine which hand leads in a bimanual task?

Right- and left-handers (n = 16 in each group) were tested on a bimanual circle task that required drawing either in the same direction (parallel) or in a mirror symmetrical coordination mode with the two hands. The authors' primary purposes were to examine the effect of circle direction on within-hand and between-hands variables and to determine whether the relation between hand lead and coordination mode (parallel or mirror symmetrical) differs for left- and right-handers. A strong relation was found between lead hand and movement condition, which depended on the direction of the movements and whether the task was parallel or mirror symmetrical. The pattern of results was similar for left- and right-handers on parallel tasks, but group differences were found with respect to mirror symmetrical tasks. At odds with the general claim that the dominant hand leads, the present results indicated that hand dominance does not generally determine which hand leads.     

The impact of perceptual, cognitive and motor factors on bimanual coordination

Bimanual coordination is governed by constraints that permit congruent movements to be performed more easily than incongruent movements. Theories concerning the origin of these constraints range from low level motor-muscle explanations to high level perceptual–cognitive ones. To elucidate the processes underlying coordinative constraints, we asked subjects to use a pair of left–right joysticks to acquire corresponding pairs of congruent and incongruent targets presented on a video monitor under task conditions designed to systematically modulate the impact of several perceptual–cognitive processes commonly required for bimanual task performance. These processes included decoding symbolic cues, detecting goal targets, conceptualizing movements in terms of goal target configuration, planning movement trajectories, producing saccades and perceiving visual feedback. Results demonstrate that constraints arise from target detection and trajectory planning processes that can occur prior to movement initiation as well as from inherent muscle properties that emerge during movement execution, and that the manifestation of these constraints can be significantly altered by the ability to visually monitor movement progress.     

Does Handedness Determine Which Hand Leads in a Bimanual Task?

Right- and left-handers (n = 16 in each group) were tested on a bimanual circle task that required drawing either in the same direction (parallel) or in a mirror symmetrical coordination mode with the two hands. The authors' primary purposes were to examine the effect of circle direction on within-hand and between-hands variables and to determine whether the relation between hand lead and coordination mode (parallel or mirror symmetrical) differs for left- and right-handers. A strong relation was found between lead hand and movement condition, which depended on the direction of the movements and whether the task was parallel or mirror symmetrical. The pattern of results was similar for left-and right-handers on parallel tasks, but group differences were found with respect to mirror symmetrical tasks. At odds with the general claim that the dominant hand leads, the present results indicated that hand dominance does not generally determine which hand leads.     

Plane of Motion Mediates the Coalition of Constraints in Rhythmic Bimanual Coordination

The authors hypothesized that the modulation of coordinative stability and accuracy caused by the coalition of egocentric (neuromuscular) and allocentric (directional) constraints varies depending on the plane of motion in which coordination patterns are performed. Participants (N = 7) produced rhythmic bimanual movements of the hands in the sagittal plane (i.e., up-and-down oscillations resulting from flexion—extension of their wrists). The timing of activation of muscle groups, direction of movements, visual feedback, and across-trial movement frequency were manipulated. Results showed that both the egocentric and the allocentric constraints modulated pattern stability and accuracy. However, the allocentric constraint played a dominant role over the egocentric. The removal of vision only slightly destabilized movements, regardless of the effects of directional and (neuro)muscular constraints. The results of the present study hint at considering the plane in which coordination is performed as a mediator of the coalition of egocentric and allocentric constraints that modulates coordinative stability of rhythmic bimanual coordination.     

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.     

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.     

Do muscles matter for coordinated action?

This article investigates coordination stability when 2 fingers of each hand periodically tap together. The main question concerns the functional origin of the symmetry tendency, which has widely been conceived as a bias toward coactivation of homologous fingers and homologous muscular portions. In Experiment 1, the symmetry tendency was independent of finger combination. In Experiment 2, virtually identical stability characteristics were revealed under full vision and no vision. In Experiment 3, symmetrical and parallel visual labels on the fingers neither stabilized nor destabilized symmetrical and parallel tapping patterns. In Experiment 4, in which the relative position of the hands was varied, it revealed that the observed stability characteristics are to be defined in a hand-centered reference frame. Because the symmetry tendency was always independent of finger combination, the authors suggest that it is not a bias toward coactivation of homologous muscle portions but instead originates on a more abstract, functional level.     

Haptic discrimination of bilateral symmetry in 2-dimensional and 3-dimensional unfamiliar displays

In five experiments, we tested the accuracy and sensitivity of the haptic system in detecting bilateral symmetry of raised-line shapes (Experiments 1 and 2) and unfamiliar 3-D objects (Experiments 3–5) under different time constraints and different modes of exploration. Touch was moderately accurate for detecting this property in raised displays. Experiment 1 showed that asymmetric judgments were systematically more accurate than were symmetric judgments with scanning by one finger. Experiment 2 confirmed the results of Experiment 1 but also showed that bimanual exploration facilitated processing of symmetric shapes without improving asymmetric detections. Bimanual exploration of 3-D objects was very accurate and significantly facilitated processing of symmetric objects under different time constraints (Experiment 3). Unimanual exploration did not differ from bimanual exploration (Experiment 4), but restricting hand movements to one enclosure reduced performance significantly (Experiment 5). Spatial reference information, signal detection measures, and hand movements in processing bilateral symmetry by touch are discussed.     

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

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

Effects of S-R and R-R compatibility on bimanual movement time

Bimanual asymmetrical movements are generally found to be slower than symmetrical movements but asymmetrical movement normally involves visual separation of targets which might account for the effect. By using a system in which the subject controls two cursors on an oscilloscope screen by moving two levers the S-R relationship on either hand can be reversed, thus providing an asymmetrical movement task without visual separation of targets. Movement times for five right-handed subjects were recorded on four unimanual and six bimanual conditions varying with respect to both S-R and R-R compatibility. In the unimanual conditions, the left hand was found to be as fast as the right when the opposite S-R relationship was used. In the bimanual tasks visual separation of targets was a relatively minor factor movement time being strongly influenced by S-R compatibility and to a lesser degree by R-R compatibility. The results suggest that compatibility, rather than being a property of a single central channel, differs, as between the two cerebral hemispheres.     

Plane of motion mediates the coalition of constraints in rhythmic bimanual coordination

The authors hypothesized that the modulation of coordinative stability and accuracy caused by the coalition of egocentric (neuromuscular) and allocentric (directional) constraints varies depending on the plane of motion in which coordination patterns are performed. Participants (N = 7) produced rhythmic bimanual movements of the hands in the sagittal plane (i.e., up-and-down oscillations resulting from flexion-extension of their wrists). The timing of activation of muscle groups, direction of movements, visual feedback, and across-trial movement frequency were manipulated. Results showed that both the egocentric and the allocentric constraints modulated pattern stability and accuracy. However, the allocentric constraint played a dominant role over the egocentric. The removal of vision only slightly destabilized movements, regardless of the effects of directional and (neuro)muscular constraints. The results of the present study hint at considering the plane in which coordination is performed as a mediator of the coalition of egocentric and allocentric constraints that modulates coordinative stability of rhythmic bimanual coordination.     

Interaction of directional,neuromuscular and egocentric constraints on the stability of preferred bimanual coordination patterns

We investigated how the relative direction of limb movements in external space (iso- and non-isodirectionality), muscular constraints (the relative timing of homologous muscle activation) and the egocentric frame of reference (moving simultaneously toward/away the longitudinal axis of the body) contribute to the stability of coordinated movements. In the first experiment, we attempted to determine the respective stability of isodirectional and non-isodirectional movements in between-persons coordination. In a second experiment, we determined the effect of the relative direction in external space, and of muscular constraints, on pattern stability during a within-person bimanual coordination task. In the third experiment we dissociated the effects on pattern stability of the muscular constraints, relative direction and egocentric frame of reference. The results showed that (1) simultaneous activation of homologous muscles resulted in more stable performance than simultaneous activation of non-homologous muscles during within-subject coordination, and that (2) isodirectional movements were more stable than non-isodirectional movements during between-persons coordination, confirming the role of the relative direction of the moving limbs in the stability of bimanual coordination. Moreover, the egocentric constraint was to some extent found distinguishable from the effect of the relative direction of the moving limbs in external space, and from the effect of the relative timing of muscle activation. In summary, the present study showed that relative direction of the moving limbs in external space and muscular constraints may interact either to stabilize or destabilize coordination patterns.     

Perception-action coupling during bimanual coordination: the role of visual perception in the coalition of constraints that govern bimanual action

Constraints pertaining to interlimb coordination have been studied extensively in the past decades. In this debate, F. Mechsner (2004) has taken a provocative position by putting primary emphasis on perceptual principles that mediate coordinative stability. Whereas the present authors agree that the role of perceptual principles is of critical importance during coordination, they take issue with Mechsner's extreme position and with the evidence forwarded to support a purely perceptual-cognitive approach to bimanual coordination. More specifically, the authors emphasize that current knowledge about brain function argues against a dualism between perception and action, criticize the presented evidence that posture manipulations during coordination provide decisive evidence against motoric and muscular constraints, and report on potential pitfalls associated with the use of visual transformation procedures to support complex coordination patterns.     

Perception—Action Coupling during Bimanual Coordination: The Role of Visual Perception in the Coalition of Constraints That Govern Bimanual Action

Constraints pertaining to interlimb coordination have been studied extensively in the past decades. In this debate, F. Mechsner (2004) has taken a provocative position by putting primary emphasis on perceptual principles that mediate coordinative stability. Whereas the present authors agree that the role of perceptual principles is of critical importance during coordination, they take issue with Mechsner's extreme position and with the evidence forwarded to support a purely perceptual-cognitive approach to bimanual coordination. More specifically, the authors emphasize that current knowledge about brain function argues against a dualism between perception and action, criticize the presented evidence that posture manipulations during coordination provide decisive evidence against motoric and muscular constraints, and report on potential pitfalls associated with the use of visual transformation procedures to support complex coordination patterns.     

Attention as a mediating variable in the dynamics of bimanual coordination

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.     

Complexity in object manipulation by Japanese macaques (Macaca fuscata): a cross-sectional analysis of manual coordination in stone handling patterns

Defined as a spontaneous stone-directed noninstrumental manipulative behavior, and comprised of multiple one-handed and (a)symmetrical/(un)coordinated two-handed patterns, stone handling (SH) is a good candidate for the study of complexity in object manipulation. We present a cross-sectional developmental analysis of SH complexity in Japanese macaques (Macaca fuscata), through the combined investigation of bimanuality, coordination, and symmetry in hand use. Bimanual SH patterns were more frequent than unimanual patterns. Among bimanual patterns, coordinated actions were more frequent than uncoordinated ones. We recorded five asymmetrical coordinated SH patterns with manual role differentiation, a form of hand use reminiscent of complex actions involving the use of tools in monkeys and apes. Bimanuality in SH was affected by body posture. Aging individuals performed less bimanual and less coordinated SH patterns than younger individuals. Our result on senescent males performing less bimanual patterns than senescent females was consistent with sex differences found in the late deterioration of complex manual movements in other species. Although some SH patterns represent a high degree of behavioral complexity, our results suggest that SH behavior is not as complex as tool-use or tool-manufacture in other nonhuman primates and hominids.     

Asymmetries in Coupling Dynamics of Perception and Action

The influence of information-based dynamics on coordination dynamics of rhythmic movement was examined with special reference to the expression of asymmetries. In Experiment 1, right-handed subjects performed unimanual, rhythmical movements in coordination with either a discrete or continuous visual display. The right hand-visual display system defined a more stable perception-action collective than the left, particularly when continuous visual information was available. In Experiment 2, the same subjects performed rhythmic bimanual movements in coordination with a continuous visual display. The action collective was inherently more stable than the perception-action collective, although similar patterns were observed at both levels. Importantly, the dynamics of the perception-action collective impinged upon the dynamics of the action collective in terms of stability. Asymmetries remained evident between limbs in the bimanual preparations, with the left hand exhibiting greater limit-cycle variability and also a tendency to more often effect transitions at the action couple. Features of dynamical models that capture characteristics of manual asymmetries are discussed.     

Effects of visual and auditory guidance on bimanual coordination complexity

Perceptual guidance of movement with simple visual or temporal information can facilitate performance of difficult coordination patterns. Guidance may override coordination constraints that usually limit stability of bimanual coordination to only in-phase and anti-phase. Movement dynamics, however, might not have the same characteristics with and without perceptual guidance. Do visual and auditory guidance produce qualitatively different dynamical organization of movement? An anti-phase wrist flexion and extension coordination task was performed under no specific perceptual guidance, under temporal guidance with a metronome, and under visual guidance with a Lissajous plot. For the time series of amplitudes, periods and relative phases, temporal correlations were measured with Detrended Fluctuation Analysis and complexity levels were measured with multiscale entropy. Temporal correlations of amplitudes and relative phases deviated from the typical 1/f variation towards more random variation under visual guidance. The same was observed for the series of periods under temporal guidance. Complexity levels for all time series were lower in visual guidance, but higher for periods under temporal guidance. Perceptual simplification of the task’s goal may produce enhancement of performance, but it is accompanied by changes in the details of movement organization that may be relevant to explain dependence and poor retention after practice under guidance.     

Kinematic parameters of hand movement during a disparate bimanual movement task in children with unilateral Cerebral Palsy

Children with unilateral Cerebral Palsy (uCP) experience problems performing tasks requiring the coordinated use of both hands (bimanual coordination; BC). Additionally, some children with uCP display involuntary symmetrical activation of the opposing hand (mirrored movements). Measures, used to investigate therapy-related improvements focus on the functionality of the affected hand during unimanual or bimanual tasks. None however specifically address spatiotemporal integration of both hands. We explored the kinematics of hand movements during a bimanual task to identify parameters of BC. Thirty-seven children (aged 10.9 ± 2.6 years, 20 male) diagnosed with uCP participated. 3D kinematic motion analysis was performed during the task requiring opening of a box with their affected- (AH) or less-affected hand (LAH), and pressing a button inside with the opposite hand. Temporal and spatial components of data were extracted and related to measures of hand function and level of impairment. Total task duration was correlated with the Jebsen–Taylor Test of Hand Function in both conditions (either hand leading with the lid-opening). Spatial accuracy of the LAH when the box was opened with their AH was correlated with outcomes on the Children’s Hand Use Experience Questionnaire. Additionally, we found a subgroup of children displaying non-symmetrical movement interference associated with greater movement overlap when their affected hand opened the box. This subgroup also demonstrated decreased use of the affected hand during bimanual tasks. Further investigation of bimanual interference, which goes beyond small scaled symmetrical mirrored movements, is needed to consider its impact on bimanual task performance following early unilateral brain injury.