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.     

Simultaneous specification of amplitudes and directions of bimanual reversal movements

Intermanual interactions are modulated by task requirements in the course of motor preparation. In particular, amplitude coupling is strong when identical amplitudes are specified concurrently for the 2 hands but relaxed when different amplitudes are specified. Similarly, directional coupling is symmetric when symmetric directions are specified concurrently but turns to parallel when parallel directions are specified. Here, the author investigated whether the modulations of amplitude coupling and directional coupling in the course of motor preparation are independent or interact. Application of the timed-response procedure, which enables one to manipulate the time available for motor preparation, in 9 participants revealed a weakly interactive pattern. Directional coupling tended to be reduced when different rather than same amplitudes were specified concurrently, and amplitude coupling tended to be reduced when parallel rather than symmetric directions were specified concurrently. In general, interactive effects were also apparent in the rates at which directions and amplitudes were specified. Those observations are consistent with the notion that intermanual amplitude and direction interference are mediated by different but partially overlapping neural structures.     

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.     

Temporal coordination during bimanual reach-to-grasp movements: The role of vision

The performance of bimanual movements involving separate objects presents an obvious challenge to the visuo-motor system: Visual feedback can only be obtained from one target at a time. To overcome this challenge overt shifts in visual attention may occur so that visual feedback from both movements may be used directly (Bingham, Hughes, & Mon-Williams, 2008; Riek, Tresilian, Mon-Williams, Coppard, & Carson, 2003). Alternatively, visual feedback from both movements may be obtained in the absence of eye movements, presumably by covert shifts in attention (Diedrichsen, Nambisan, Kennerley, & Ivry, 2004). Given that the quality of information falls with increasing distance from the fixated point, can we obtain the level of information required to accurately guide each hand for precision grasping of separate objects without moving our eyes to fixate each target separately? The purpose of the current study was to examine how the temporal coordination between the upper limbs is affected by the quality of visual information available during the performance of a bimanual task. A total of 11 participants performed congruent and incongruent movements towards near and/or far objects. Movements were performed in natural, fixate-centre, fixate-left, and fixate-right vision conditions. Analyses revealed that the transport phase of incongruent movements was similar across vision conditions for the temporal aspects of both the transport and grasp, whereas the spatial aspects of grasp formation were influenced by the quality of visual feedback. We suggest that bimanual coordination of the temporal aspects of reach-to-grasp movements are not influenced solely by overt shifts in visual attention but instead are influenced by a combination of factors in a task-constrained way.     

Response-response compatibility during bimanual movements: Evidence for the conceptual coding of action

The present study investigated response-response (R-R) compatibility in a bimanual keypressing task. Numeric and spatial stimuli were used to cue responses for each hand. Two groups of participants differed in terms of the stimulus-response mappings for the numeric stimuli. For one group, the numeric stimuli were mapped so that the same number for each hand indicated responses that were anatomically compatible (e.g., index finger of both hands). For the other group, the same number for each hand indicated responses that were left-right compatible (e.g., leftmost finger of both hands). The spatial stimuli were mapped in a spatially compatible manner to the responses for both groups. For numeric stimuli, reaction times (RTs) were faster when the same number indicated the response for each hand, regardless of the mapping. For the spatial stimuli, RTs were determined not only by the pairing of stimuli or responses, but also by how the responses were indicated by numeric stimuli. The results indicate that R-R compatibility effects are mediated by abstract codes that reflect individuals’ conceptualizations of their actions.     

Oculomotor interference during manual response preparation: evidence from the response-cueing paradigm

Preparation provided by visual location cues is known to speed up behavior. However, the role of concurrent saccades in response to visual cues remains unclear. In this study, participants performed a spatial precueing task by pressing one of four response keys with one of four fingers (two of each hand) while eye movements were monitored. Prior to the stimulus, we presented a neutral cue (baseline), a hand cue (corresponding to left vs. right positions), or a finger cue (corresponding to inner vs. outer positions). Participants either remained fixated on a central fixation point or moved their eyes freely. The results demonstrated that saccades during the cueing interval altered the pattern of cueing effects. Finger cueing trials in which saccades were spatially incompatible (vs. compatible) with the subsequently required manual response exhibited slower manual RTs. We propose that interference between saccades and manual responses affects manual motor preparation.     

Perceived movement of the afterimage during eye movements

The question of whether an afterimage viewed in a dark field appears to move during eye movement was studied by comparing recordings of eye movements with recordings of reports of perceived movement. The correlation was found to be quite good even under conditions where the eye movements were spontaneous rather than specifically directed. The results were taken to support the hypothesis that the behavior of the retinal image is “interpreted” by taking into account information concerning what the eyes are doing.     

The preparation of reach-to-grasp movements in adults, children, and children with movement problems

This study explored the use of advance information in the control of reach-to-grasp movements. The paradigm required participants to reach and grasp illuminated blocks with their right hand. Four target blocks were positioned on a table surface, two each side of the mid-saggital plane. In the complete precue condition, advance information precisely specified target location. In the partial precue condition, advance information indicated target location relative to the midsaggital plane (left or right). In the null condition, the advance information was entirely ambiguous. Participants produced fastest responses in the complete precue condition, intermediate response times in the partial condition, and the slowest responses in the null condition. This result was observed in adults and four groups of children including a group aged 4-6 years. In contrast, children with Developmental Coordination Disorder (DCD, n = 11, aged 7-13 years) showed no advantage of partial precueing. Movement duration was determined by target location but was unaffected by precue condition. Movement duration was a clear function of age apart from children in the DCD group who showed equivalent movement times to those of the youngest children. These findings provide important insights into the control of reach-to-grasp movements and highlight that partial cues are exploited by children as young as 4 years but are not used in situations of abnormal development.     

Effect of starting position on reproduction of movement: further evidence of interference between location and distance information

Two experiments done with a short-term memory paradigm examined the influence of shifts in the starting position on the reproduction of kinesthetic location (Exp. 1) and on distance cues (Exp. 2). We assessed possible causes of the systematic pattern of undershooting and overshooting as related to the shift in the starting position. In each experiment, two groups of 10 students were given 25 trials, and each had criterion and reproduction tasks involving linear-positioning movements with a 10-sec. retention interval. Each experiment had two independent variables, the group of subjects and the shift in the starting position. The two groups differed in the possible sources of information, the distance moved (Exp. 1) or the end-location (Exp. 2), which were assumed to cause undershooting and overshooting during reproduction. Analysis showed that the information about the distance moved may produce undershooting and overshooting in reproduction of the end-location (Exp. 2). Also, the information about the end-location may produce undershooting and overshooting in reproduction of the distance moved (Exp. 2). The findings were further evidence of interference between location and distance cues in motor short-term memory.     

Probing the motor program: Effects of output competition during movement preparation

A probe reaction-time paradigm was used to investigate the capacity demands of planning rapid aiming movements. Subjects were required to respond either vocally or manually to an auditory probe presented during the reaction-time interval preceding a pointing response. When a vocal response was required probe reaction time increased systematically with the complexity of the pointing movements. Presumably this is because a more complex task requires more programming resources. When a manual response was required, reaction-time data for both the pointing task and the probe indicate that the structural constraints inherent in programming two similar movements may force subjects to engage in common response preparation. The methodological and theoretical implications of these findings are discussed.     

Dissociation of covert and overt spatial attention during prehension movements: Selective interference effects

In four experiments, the influence of distractor objects on the temporal evolution of the reach-tograsp movement toward a target object (an apple) was examined. In the first experiment, the distractor was another apple, which moved laterally behind the target and occasionally changed direction toward the target, thus becoming the to-be-grasped object. In the second and third experiments, the distractor was a stationary piece of fruit, which sometimes became the to-be-grasped object because of a change in illumination. The fourth experiment was a combination of the first two experiments. In all cases, selective interference effects on the transport and manipulation components were observed only when attention to the distractor was covert rather than overt. It is proposed that covert visuospatial attention selects information about distracting but potentially important stimuli, such that a registration of significance is accomplished without the need to process all available information.     

Integration of posture and movement: contributions of Sherrington, Hess, and Bernstein

Neural mechanisms that integrate posture with movement are widespread throughout the central nervous system (CNS), and they are recruited in patterns that are both task- and context-dependent. Scientists from several countries who were born in the 19th century provided essential groundwork for these modern-day concepts. Here, the focus is on three of this group with each selected for a somewhat different reason. Charles Sherrington (1857-1952) had innumerable contributions that were certainly needed in the subsequent study of posture and movement: inhibition as an active coordinative mechanism, the functional anatomy of spinal cord-muscle connectivity, and helping set the stage for modern work on the sensorimotor cortex and the corticospinal tract. Sadly, however, by not championing the work of his trainee and collaborator, Thomas Graham Brown (1882-1965), he delayed progress on two key motor control mechanisms: central programming and pattern generation. Walter Hess (1881-1973), a self-taught experimentalist, is now best known for his work on CNS coordination of autonomic (visceral) and emotional behavior. His contributions to posture and movement, however, were also far-reaching: the coordination of eye movements and integration of goal-directed and "framework" (anticipatory set) motor behavior. Nikolai Bernstein (1896-1966), the quintessence of an interdisciplinary, self-taught movement neuroscientist, made far-reaching contributions that were barely recognized by Western workers prior to the 1960s. Today, he is widely praised for showing that the CNS's hierarchy of control mechanisms for posture and movement is organized hand-in-hand with distributed and parallel processing, with all three subject to evolutionary pressures. He also made important observations, like those of several previous workers, on the goal focus of voluntary movements. The contributions of Sherrington, Hess, and Bernstein are enduring. They prompt thought on the philosophical axioms that appear to have driven their research, and the continual need for emphasis on interdisciplinary, comparative, and transnational approaches to advance movement neuroscience.     

Induced rotary movement during eye movements: displays with unequal spacing of pattern

In my previous (Reinhardt-Rutland, 1982) study, I suggested that eye movements enhance induced rotary movement. However, low salience of absolute displacement might also explain the results, as displays were covered with large numbers of equally spaced radial pattern elements. To test these competing hypotheses in the present study, I used an unequally spaced pattern in two displays. Common to each display was an annulus: In one display, the common annulus surrounded a disk, and in the other display the common annulus was surrounded by another annulus. In any trial, one component rotated and the other was stationary while for 40 s the subject's eyes followed a circular path concentric with the display; subjects timed those occasions when perceived stronger rotation resided in the common annulus. Despite an unequally spaced pattern, absolute displacement had a barely significant effect. Instead, perceived stronger rotation mostly resided in a display's more central component. I concluded therefore that eye movements enhance induced rotary movement.     

Space-time behavior of single and bimanual rhythmical movements: data and limit cycle model

How do space and time relate in rhythmical tasks that require the limbs to move singly or together in various modes of coordination? And what kind of minimal theoretical model could account for the observed data? Earlier findings for human cyclical movements were consistent with a nonlinear, limit cycle oscillator model (Kelso, Holt, Rubin, & Kugler, 1981) although no detailed modeling was performed at that time. In the present study, kinematic data were sampled at 200 samples/second, and a detailed analysis of movement amplitude, frequency, peak velocity, and relative phase (for the bimanual modes, in phase and antiphase) was performed. As frequency was scaled from 1 to 6 Hz (in steps of 1 Hz) using a pacing metronome, amplitude dropped inversely and peak velocity increased. Within a frequency condition, the movement's amplitude scaled directly with its peak velocity. These diverse kinematic behaviors were modeled explicitly in terms of low-dimensional (nonlinear) dissipative dynamics, with linear stiffness as the only control parameter. Data and model are shown to compare favorably. The abstract, dynamical model offers a unified treatment of a number of fundamental aspects of movement coordination and control.     

Brain injury and movement recall: Preselection, active-passive and interference effects

Stroke patients with unilateral lesions were compared with age-controls and students on their ability to reproduce a terminal location established kinesthetically by a previous movement. Conditions for the criterion movement differed over active/passive and preselected/constrained (experiment 1) and whether the retention interval between the criterion and recall movements involved mental rehearsal of the criterion movement or yes/no responding to a mental arithmetic task (experiment 2). Whereas students showed more accurate recall with little effect of criterion movement condition, patient groups showed a preselection effect, but only with active movements. A preferred hand advantage observed for the patient controls did not occur with stroke patients, and prevention of mental rehearsal during the retention interval disrupted recall more for the stroke patients. These findings are interpreted in terms of hemisphere-specific coding strategies whose relative use depends on the attentional demands of the task.     

A new approach to cerebral asymmetry: RT differences in simultaneous bimanual finger movement during verbal and nonverbal tasks

This study was aimed at testing a new approach for examination of functional laterality based on hemispheric specialization. The subjects had to perform verbal (words/nonwords) and nonverbal (similar/different patterns) discrimination. The separation of the two hemispheres during information processing was realized by requiring a simultaneous response of both index fingers. The obtained over-all reaction times (RT) were faster for verbal than for pattern tasks. Considering the RTs for solely the particular, faster response of one or the other index finger, the right index finger turned out to be faster on verbal tasks whereas the left one dominated on pattern tasks. According to the hypothesis that the faster hand indicates the more active (contralateral) hemisphere, it can be assumed that words are responded to more quickly when processed in the left hemisphere. On the other hand, patterns are responded to more quickly when the right hemisphere is active. These results suggest that each hemisphere may be capable of processing verbal and nonverbal material; the speed of information processing, however, is faster in the more adept one.     

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.