Anisotropic tracking: evidence for automatic synergy formation in a bimanual task

Investigation of interlimb synergy has become synonymous with the study of coordination dynamics and is largely confined to periodic movement. Based on a computational approach this paper demonstrates a method of investigating the formation of a novel synergy in the context of stochastic, spatially asymmetric movements. Nine right-handed participants performed a two degrees of freedom (2D) "etch-a-sketch" tracking task where the right hand controlled the horizontal position of the response cursor on the display while the left hand controlled the vertical position. In a pre-practice 2D tracking task, measures of phase lag between the irregularly moving target and the response showed that participants controlled left and right hands independently, performance of the right hand being slightly superior to the left. Participants then undertook 4 h 16 min distributed practice of a one degree of freedom etch-a-sketch task where the target was constrained to move irregularly in only the 45 degrees direction on the display. To track such a target accurately participants had to make in-phase coupled stochastic movements of the hands. In a post-practice 2D task, measures of phase lag showed anisotropic improvement in performance, the amount of improvement depending on the direction of motion on the display. Improvement was greatest in the practised 45 degrees and least in the orthogonal 135 degrees direction. Best and worst performances were no longer in the directions associated with right and left hands independently, but in directions requiring coupled movements of the two hands. These data support the proposal that the nervous system can establish a model of novel coupling between the hands and thereby form a task-dependent bimanual synergy for controlling the stochastic coupled movements as an entity.     

Sex Differences in Perception: Exploring the Integration of Sensory Information with Respect to Vision and Proprioception

Inter- and intra-modal matching by right hand preferred Norwegian adults were tested using a matching task. The task required them to locate target pins visually, proprioceptively or in combination, while matching was always carried out without vision. When combined scores for both hands were analysed, the females showed superior performance in the intra-modal matching condition. For both groups, when the scores for the two hands were treated separately, superior performance was obtained in using the right hand to match under the visual condition. The only significant sex difference relative to hand preference was that the females matched the target more accurately with the right hand in the intra-modal matching condition. These findings are discussed relative to inter-hemispheric processing     

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.     

A developmental analysis of the relationship between hand preference and performance: I. Preferential reaching into hemispace

The present study describes a developmental performance measure of hand preference that considers task complexity and position in hemispace. Eighty right-handed children and adults (ages 3-4, 6-7, 9-10, 18-24) were observed for hand selection responses to 2 unimanual tasks (simple vs complex) across positions in hemispace. Results revealed an age-related trend in the tendency to use the preferred hand in right and left hemispace. While the adult's and 3- to 4-year-old's preferred hand use decreased as they moved into left hemispace, children between the ages of 6 and 10 years tended to use their preferred hands consistently throughout both regions of hemispace. The relationship between hand preference and skilled, cost-efficient performance throughout development are discussed.     

Effects of S-R Compatibility and Task Difficulty on Unimaimual Movement Time

Using a system in which S controlled a cursor on an oscilloscope screen by moving a lever, the S-R relationship on either hand could be reversed. This system was used in two experiments designed to investigate the effect on unimanual movement time of varying S-R compatibility, and task difficulty as defined by Fitts (1954). The results indicated the necessity of specifying task difficulty in an investigation of S-R compatibility. Further, it was found that as task difficulty increased the difference in performance between the right and left hands became more marked.     

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.     

Hand dominance, attention, and the choice between responses

Thirty pairs of matched right- and left-handed subjects carried out a serial C.RT task involving reaches between 2.4 cm square contact grids set at 2.4 cm intervals in a horizontal line. In one condition they responded with the right hand alone, in a second with the left and in a third they had to choose between hands on each trial responding to left side grids with the left hand and to right side grids with the right.

All subjects took longer to respond when they had to choose between hands than when they used either hand alone. In the one-hand conditions neither group showed any effect of hand dominance. When choices had to be made between hands both groups responded faster with their dominant than with their nondominant hands.

Analysis of the results was undertaken in terms of two hypothetical systems of instructions which might control machines designed to carry out similar tasks. It is concluded that in some tasks hand dominance can be described in terms of an attentional bias towards the field of operation of one effector rather than another. In the present task the effects of such possible attentional biases were shown not to interact with other effects, such as the facilitation of successive responses by repeated use of the same limb. These latter effects seem to depend on processes underlying execution of responses rather than choices between responding limbs.     

Prism adaptation in alternately exposed hands

We assessed intermanual transfer of the proprioceptive realignment aftereffects of prism adaptation in right-handers by examining alternate target pointing with the two hands for 40 successive trials, 20 with each hand. Adaptation for the right hand was not different as a function of exposure sequence order or postexposure test order, in contrast with adaptation for the left hand. Adaptation was greater for the left hand when the right hand started the alternate pointing than when the sequence of target-pointing movements started with the left hand. Also, the largest left-hand adaptation appeared when that hand was tested first after exposure. Terminal error during exposure varied in cycles for the two hands, converging on zero when the right hand led, but no difference appeared between the two hands when the left hand led. These results suggest that transfer of proprioceptive realignment occurs from the right to the left hand during both exposure and postexposure testing. Such transfer reflects the process of maintaining spatial alignment between the two hands. Normally, the left hand appears to be calibrated with the right-hand spatial map, and when the two hands are misaligned, the left-hand spatial map is realigned with the right-hand spatial map.     

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. Eleven 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.     

Hand preference in children with developmental coordination disorders: cause and effect?

Inter- and intra-modal matching by eight-year-old children diagnosed as having hand-eye coordination problems (HECP) and categorized as left-handed, together with a left-handed control group of children without such problems, were tested using a manual sensory matching task. The task required the children to locate target pins, visually (seen target), proprioceptively (felt target) or in combination (felt and seen target), while matching to the located target was always carried out without vision. Performance was superior when the target was located visually or visually/proprioceptively for both groups of children. These results question the conclusion that intra-modal will always be more accurate than inter-modal matching. When the combined scores for both hands were analyzed, the HECP children showed inferior performance to the control children in both inter- and intra-modal matching. Separate right and left hand analyses, demonstrated that the differences between the HECP group and control children could be accounted for by lowered performances when the right hand (nonpreferred) was used to match the located target position. Putative neurological disorders related to the development of the hemisphere controlling the nonpreferred hand (left hemisphere) are invoked to account for the poor performance with the nonpreferred hand of the HECP children.     

Cerebral Organization of Motor Programming and Verbal Processing as a Function of Degree of Hand Preference and Familial Sinistrality

Seventy-six right- and left-handed subjects responded to monaurally presented verbal stimuli (CVs) using their right and left hands on separate occasions. Both degree of hand preference and familial sinistrality (FS) were taken into account. It was found that, contrary to expectation, the manual response interfered with the verbal perception task, but only in the consistent strong handers. The pattern of interference suggests that those with a consistent hand preference (right or left) have general motor programming in the left hemisphere. Those with an inconsistent strong hand preference probably have some degree of general motor programming in both hemispheres. No effect for FS was found for the lateralization of verbal processing or general motor programming.     

Performance Asymmetries in Computer Mouse Control of Right-Handers, and Left-Handers with Left- and Right-Handed Mouse Experience

Precision and general computer mouse aiming performance by right-handers (RH-RM) and left-handers with right-handed mouse experience (LH-RM) and by left-handers with left-handed mouse experience (LH-LM) were compared. A number of performance measures, such as reaction time, time to reach a target, time to click on target, and cursor trajectory, were analyzed. Superficially, specific hand experience seemed to dictate performance asymmetries, but a closer look revealed interactions between hand preference and hand performance. That finding has implications for theories of handedness. In addition, precision and general directional aiming with the mouse cursor showed a clear right hand superiority in reaction time in both RH-RM and LH-RM subjects, whereas LH-LM subjects showed no lateral asymmetries. Finally, the overall time taken for the task, averaged across groups and conditions, favored the experienced hand by some 180 ms. In practical terms, that is not a large difference, especially because the difference will be reduced with practice. Thus, the use of the inexperienced hand can be advocated when there is a need to forestall or ameliorate repetitive stress in the experienced hand.     

Complexity matching and coordination in individual and dyadic performance

Complexity matching is a measure of coordination based on information exchange between complex networks. To date, studies have focused mainly on interpersonal coordination, but complexity matching may generalize to interacting networks within individuals. The present study examined complexity matching in a double, coordinated Fitts' perceptual-motor task with comparable individual and dyadic conditions. Participants alternated touching targets with their left and right hands in the individual condition, or analogously with the left hand of one partner and the right hand of another in the dyadic condition. In Experiment 1, response coupling was manipulated by making targets drift either randomly or contingently based on prior responses. Here, drift refers to the variability in the target movements between response locations. Long-range correlations in time series of inter-response intervals exhibited complexity matching between the left and right hands of dyads and individuals. Response coupling was necessary for complexity matching in dyads but not individuals. When response coupling was absent in the dyadic condition, the degree of complexity matching was significantly reduced. Experiment 2 showed that the effect of coupling was due to interactions between left and right responses. Results also showed a weak, negative relationship between complexity matching and performance as measured by total response time. In conclusion, principles and measures of complexity matching apply similarly within and between individuals, and perceptual-motor performance can be facilitated by loose response coupling.     

Frames of reference in perceptual-motor learning: Evidence from a blind manual positioning task

Participants moved a joystick to bring a computer-displayed cursor to each of six on-screen target locations arrayed around the center of the screen. At the start of each trial, the stick rested vertically, with a cursor occupying the center of the screen. A target appeared at another location and as soon as the stick was moved away from its rest position the cursor disappeared until the participant pressed a trigger on the stick to indicate when s/he thought the stick-controlled cursor was at the target site. With training, participants improved on the blind positioning task, but when conditions changed their performance suffered. Changing the hand used in the task or the location of the stick caused approximately equal disruptions, but changing both hand and location was significantly more disruptive than changing just one feature. The results support the hypothesis that perceptual-motor learning entails coding of extrinsic (spatial coordinates) as well as intrinsic (postural or body movement) information. Received: 14 October 1999 / Accepted: 6 November 2000     

Hemispace and information control by the two cerebral hemispheres: which interaction?

Two experiments employing subjects with different experience in tactile discrimination (blind and seeing subjects) were carried out to investigate the effect of the space location of stimuli on the information processing activity of the two cerebral hemispheres. An angle discrimination task that yields a right hemisphere superiority was used. In Experiment 1, seeing subjects showed a general superiority of the left hand (right hemisphere) which was more pronounced in the left hemispace with respect to the central and the right hemispace performance. In Experiment 2, blind subjects showed a significant superiority of the left hand in the central and in the left hemispace and no difference between the two hands in the right hemispace. In both experiments hemispace differences were due only to the modification of the left hand (right hemisphere) performance. These results suggest that the hemispace control by the contralateral hemisphere interacts only with the activity of the hemisphere dominant in the information processing.     

Hand preference in simultaneous unimanual tasks: a preliminary examination

The aim of the current investigation was to determine the pattern of hand use during simultaneous unimanual tasks. Two studies were conducted. The first experiment examined the pattern of hand use in a catching task, while performing a secondary writing task. Results showed that individuals had a decreased tendency to catch with their preferred hand when their preferred hand was occupied, in comparison to when the preferred hand was unoccupied. The second experiment examined the pattern of hand use during a support and reach task, where the use of both hands was required. Here, results indicated that participants preferred to support themselves with their nonpreferred hand and reach with preferred hand toward right hemispace. With respect to left hemispace, participants showed the reverse pattern. This pattern of hand use indicates an important role for the nonpreferred hand, which has been relatively unexplored by researchers.     

Manual localization of lateralized visual targets

The effects of visual field, responding limb and extrapersonal space on the ability to localize visual targets using slow positioning movements of the arm were examined. Special contact lenses were used to lateralize visual information and to make comparisons with localization under monocular control conditions. Subjects made slow positioning movements to place a cursor directly beneath target lights. They saw target lights but not the moving limb during the trial. For directional error, results indicated that subjects were more accurate localizing targets lateralized to the right hemisphere than targets lateralized to the left hemisphere, indicating right hemisphere superiority for localization of visual targets in grasping space. Localization performance was significantly better with the right hand than the left hand. the left hand demonstrated a directional bias to the right of the targets. Responding hand and visual field did not interact. Finally, contrary to subjects' awareness and verbal reports, target localization was not less accurate in lens than in monocular control conditions. This was true for both amplitude and directional error. This is consistent with other studies where visual information about limb position is not available.     

Selective attention and cerebral dominance in perceiving and responding to speech messages

The effect of attention on cerebral dominance and the asymmetry between left and right ears was investigated using a selective listening task. Right handed subjects were presented with simultaneous dichotic speech messages; they shadowed one message in either the right or left ear and at the same time tapped with either the right or the left hand when they heard a specified target word in either message. The ear asymmetry was shown only when subjects' attention was focused on some other aspect of the task: they tapped to more targets in the right ear, but only when these came in the non-shadowed message; they made more shadowing errors with the left ear message, but chiefly for non-target words. The verbal response of shadowing showed the right ear dominance more clearly than the manual response of tapping. Tapping with the left hand interfered more with shadowing than tapping with the right hand, but there was little correlation between the degree of hand and of ear asymmetry over individual subjects. The results support the idea that the right ear dominance is primarily a quantitative difference in the distribution of attention to left and right ear inputs reaching the left hemisphere speech areas. This affects both the efficiency of speech perception and the degree of response competition between simultaneous verbal and manual responses.     

Effect of slow movement execution on cognitive function

We investigated the effect of slow paced movement on cognitive function. The task movement was a dual-task performance composed of a continuous forearm rotation for the right hand and a simple reaction task for the left hand. Exp. 1 was designed to compare reaction time during performance at a slow pace to that at medium pace by 14 female undergraduate students. The mean reaction time for the left hand under the Slow Pace was significantly longer than that under the Middle Pace condition (p < .05), which showed that the subjects were required to give more attention to right-hand performance at the slow pace as it was difficult. Exp. 2 examined changes in reaction time when using the left hand that were associated with the learning of a slow paced task while using the right hand. Twenty-three female undergraduate students participated and repeated the task 6 times. The 3 sec. prior to and the 3 sec. after each auditory stimulus were used to establish rotation speed and mean coefficients of variation. The mean coefficients of variation, evaluated as within-subject variability, showed a significantly positive correlation with reaction time at Trials 1 and 6 for prestimulus and Trials 5 and 6 for poststimulus. Over successive trials participants continued performing the primary forearm task at a constant slow pace before and after receiving auditory stimuli, and this progress was related to a decrease in reaction time. Further, the sense of concentration evaluated by the subjects poststimulus was significantly higher than that prestimulus (p < .01). Performance at a constant speed, which was much slower than the ordinary or preferred speed of each subject, may have had a strong effect on their ability to remain conscious of movement execution.     

Visual line bisection in sinistrals and dextrals as a function of hemispace, hand, and scan direction

Visual line bisection was investigated in 26 sinistral and 24 dextral subjects as a function of hemispace, hand and scan direction. An ANOVA revealed significant main effects for hand preference, due to the mean bisection errors of dextral subjects being significantly leftward of those of sinistral subjects; for hand, due to the bisection errors of the left hand being significantly to the left of the right hand; and for scan, due to the bisection errors following a left scan being significantly to the left of a right scan. One significant interaction was found, that between hand and direction of scan, due to a significant difference between left and right hands following a scan from the left but not following a scan from the right. For dextral subjects the leftward bisection errors of the left and right hands following a scan from the left, but not for a scan from the right, differed significantly from the midpoint. For sinistral subjects the leftward bisection errors following a scan from the left and rightward bisection errors following a scan from the right differed significantly from the midpoint for the left hand but not for the right hand. No significant main effect or interactions for hemispace were found. This confirms that both sinistral and dextral subjects display pseudoneglect when using their preferred hand and scanning from the left. However, sinistrals, but not dextrals, will display reversed pseudoneglect when using their preferred hand and adopting a scan direction from the right. These results are discussed in terms of the interaction between three factors, whose influence can jointly and severally produce misbisections, hemispheric specialisation for visuospatial function, hemispheric activation for a manual response, and the allocation of visual attention.