Right-handers' reaching in contralateral hemispace: a kinematic observation

C. Gabbard and C. Helbig (2004) found, when examining seated participants' limb selection for reaching and grasping in hemispace, that right-handers preferred to switch to the nondominant left arm for objects located approximately 20 degrees horizontally from body midline (90 degrees) in left hemispace. In the present study, the authors examined 13 strongly lateralized seated right-handers' kinematics of reaching to object positions ranging from body midline to 40 degrees horizontally in left hemispace. Participants executed faster reaches with the left arm than with the right arm to objects placed 20 degrees-40 degrees from midline, whereas they did not change the proportion of time they spent accelerating the hands when the position of the object changed. A 2nd main finding was an increase in the left hand's trajectory curvature as object position moved farther from midline, with a corresponding decrease in the contribution of upper-arm motion to the reach. Those observations suggest that the switch from dominant right-arm reaching to nondominant left-arm reaching in left hemispace reported in the aforementioned study may have emerged from a shift from a shoulder-driven reach to an elbow-driven action.     

Spatial compatibility effects on the same side of the body midline

Stimulus-response compatibility effects have been hypothesized to result (a) from a subject's innate tendency to respond in the direction of the source of stimulation, (b) from a correspondence between the spatial codes associated with the effector and the stimulus, or (c) from an attentional bias favoring the effector located in the same hemispace as the command signal. Two experiments were conducted to test these three hypotheses. In Experiment 1 the subjects were requested to make unimanual discriminative key-pressing responses to two light stimuli, both appearing to either the right or left of the fixation point. In one condition the two hands were in anatomical position (uncrossed); in the other they were crossed. The procedure of Experiment 2 was similar to that of Experiment 1 with the exception that both hands, always in an uncrossed position, were placed on the same side of the body midline (on the right or left). The results showed that the compatibility effect depends on a correspondence between the spatial codes associated with the location of the effector and the location of the command stimulus.     

Effects of Expectancy and Attention in Vibrotactile Choice Reaction Time Tasks

The orienting of attention in space has not been considered in the tactile domain. This issue is examined using a modified version of a visual paradigm initially adopted by Posner, Snyder, and Davidson (1980), which manipulates the probability of a stimulus occurring at different spatial locations. Slower RTs at an unexpected stimulus location are thought to reflect the time required to shift attention from the expected to the unexpected location. In two experiments involving vibrotactile choice RT between left and right hands, the two hands were either crossed or uncrossed, and the hands were held both on the left side of the body, both on the right, or one on either side of the midline. There was no evidence to suggest that spatial location (left or right) affected the orienting of attention in the tactual modality. As predicted, RTs were slower when the arms were crossed compared with uncrossed, though this effect was smaller for the expected trials. A coding conflict hypothesis may explain both these findings, but the smaller effect in the expected trials may also reflect attentional factors. Both the relative and absolute location of the hands affected the magnitude of the crossed-arm effect and indicated that attention may play a role in the perceptual division of space into left and right sides. Possible reasons for hand or hemispace asymmetries in different simple and choice RT paradigms were discussed.     

Unilateral hemispheric activation does not affect free-viewing perceptual asymmetries

Strong leftward perceptual biases have been reported for the selection of the darker of two left/right mirror-reversed luminance gradients under free-viewing conditions. This study investigated the effect of unilateral hemispheric activation on this leftward bias in two groups of dextrals (N = 52 and N = 24). In Experiment 1, activation was manipulated by asking participants to tap with their left or right fingers along their midline. In Experiment 2, participants clenched their left or right hands in their respective hemispaces. Participants selected the stimulus that was darker on the left-hand side 73% of the time. Despite manipulations of activation strength and hemispace, activation had no effect on the asymmetry. If activation was important, the leftward bias should have been enhanced when the left hand/right hemisphere was active and reduced (or reversed) when the right hand/left hemisphere was active. The contribution of left-to-right scanning biases to free-viewing perceptual asymmetries is discussed as an alternative.     

Selective spatial attention and length representation in normal subjects and in patients with unilateral spatial neglect

The aim of this study was to assess whether perceptual representation along the horizontal axis is affected by hemispace position of the stimulus or by orienting attention to one side. Ten control subjects and 10 right brain damaged patients with left unilateral spatial neglect (USN) were asked to bisect lines of five lengths in three space positions (left, center, right) and under three cueing conditions (no cue, left cue, right cue). Normal controls showed significant displacement of bisection opposite to the side of hemispace presentation and toward the side of cueing. USN patients showed a bisection error toward the right end which increased with lines placed in the left hemispace and decreased with lines placed in the right hemispace and when attention was oriented toward the left side. We conclude that (1) In absence of cues normal subjects tend to overestimate the portions of space closer to their body midline; (2) both normal and USN patients tend to overestimate portions of space that they direct their attention to; (3) USN patients' performance without cueing is consistent with an attentional shift toward the right hemispace implying a gradient of overestimation of the right-most portions of space. A common neural substratum for directing attention and space representation can explain these findings.     

Mapping of extracorporeal space by vibrotactile reaction times: a far-left-side disadvantage

Vibrotactile reaction times in normal dextrals were measured for the two hands separately when either hand was located at each of seven possible positions: 90 degrees, 45 degrees, and 15 degrees to the left and right of the chest midline, and at the midline itself (0 degrees). Reaction times for the two hands did not differ and there was no Hand by Position interaction. At 90 degrees left, reaction times were significantly slower than at any other position except 45 degrees right. However, none of the other positions, including 45 degrees right, differed from each other. Performance in this task, therefore, was relatively uniform from 90 degrees right to 45 degrees left, but markedly slower at 90 degrees left. This far-left-side disadvantage may reflect a difficulty (for dextrals) in focussing covert attention in the far-left part of space for a block of trials. Since vibrotactile reaction times are sensitive to attentional factors in normal subjects, the paradigm should allow quantification of the clinical symptoms of the hemineglect syndrome; some preliminary observations of this syndrome with another vibrotactile design are reported.     

Sensory and motor aspects of pseudoneglect, hemifield, and hemispace in the tactile modality

Two experiments were conducted with right-handed adult subjects to investigate motor and sensory components of a tactual line bisection task performed under three conditions: at midline, in the left, and in the right hemispaces. In the sensory experiment we found a left-hand rather than a right-hand superiority under the midline condition and, in the motor experiment, a right-hand rather than a left-hand superiority. The results were discussed with respect to hemispheric specialization and hemispace theories. Furthermore, we found a pseudoneglect (subjects bisected to the left of the midpoint) in the sensory experiment and a surprising reversed pseudoneglect (subjects bisected to the right of the midpoint) in the motor experiment.     

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.     

Where you look can influence haptic object recognition

We investigated whether the relative position of objects and the body would influence haptic recognition. People felt objects on the right or left side of their body midline, using their right hand. Their head was turned towards or away from the object, and they could not see their hands or the object. People were better at naming 2-D raised line drawings and 3-D small-scale models of objects and also real, everyday objects when they looked towards them. However, this head-towards benefit was reliable only when their right hand crossed their body midline to feel objects on their left side. Thus, haptic object recognition was influenced by people's head position, although vision of their hand and the object was blocked. This benefit of turning the head towards the object being explored suggests that proprioceptive and haptic inputs are remapped into an external coordinate system and that this remapping is harder when the body is in an unusual position (with the hand crossing the body midline and the head turned away from the hand). The results indicate that haptic processes align sensory inputs from the hand and head even though either hand-centered or object-centered coordinate systems should suffice for haptic object recognition.     

Constraints on Arm Selection Processes When Reaching: Degrees of Freedom and Joint Amplitudes Interact to Influence Limb Selection

With an interest in identifying the variables that constrain arm choice when reaching, the authors had 11 right-handed participants perform free-choice and assigned-limb reaches at 9 object positions. The right arm was freely selected 100% of the time when reaching to positions at 30° and 40° into right hemispace. However, the left arm was freely selected to reach to positions at ?30° and ?40° in left hemispace 85% of the time. A comparison between free- and assigned-limb reaching kinematics revealed that free limb selection when reaching to the farthest positions was constrained by joint amplitude requirements and the time devoted to limb deceleration. Differences between free- and assigned-arm reaches were not evident when reaching to the midline and positions of ±10°, even though the right arm was freely selected most often for these positions. Different factors contribute to limb selection as a function of distance into a specific hemispace.     

Constraints on arm selection processes when reaching: degrees of freedom and joint amplitudes interact to influence limb selection

With an interest in identifying the variables that constrain arm choice when reaching, the authors had 11 right-handed participants perform free-choice and assigned-limb reaches at 9 object positions. The right arm was freely selected 100% of the time when reaching to positions at 30° and 40° into right hemispace. However, the left arm was freely selected to reach to positions at -30° and -40° in left hemispace 85% of the time. A comparison between free- and assigned-limb reaching kinematics revealed that free limb selection when reaching to the farthest positions was constrained by joint amplitude requirements and the time devoted to limb deceleration. Differences between free- and assigned-arm reaches were not evident when reaching to the midline and positions of ±10°, even though the right arm was freely selected most often for these positions. Different factors contribute to limb selection as a function of distance into a specific hemispace.     

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.     

Failure to remap visuotactile space across the midline in the split-brain.

We examined the effect of posture change on the representation of visuotactile space in a split-brain patient using a cross-modal congruency task. Split-brain patient J.W. made speeded elevation discrimination responses (up versus down) to a series of tactile targets presented to the index finger or thumb of his right hand. We report congruency effects elicited by irrelevant visual distractors placed either close to, or far from, the stimulated hand. These cross-modal congruency effects followed the right hand as it moved within the right hemispace, but failed to do so when the hand crossed the midline into left hemispace. These results support recent claims that interhemispheric connections are required to maintain an accurate representation of visuotactile space.     

Effects of response eccentricity and relative position on orthogonal stimulus-response compatibility with joystick and keypress responses

When unimanual left-right movement responses are made to up-down stimuli, performance is better with the up-right/down-left mapping when responding in the right hemispace and with the up-left/down-right mapping when responding in the left hemispace. We evaluated whether this response eccentricity effect is explained best in terms of rotational properties of the hand (the end-state comfort hypothesis) or asymmetric coding of the stimulus and response alternatives (the salient features coding hypothesis). Experiment 1 showed that bimanual keypresses yield a response eccentricity effect similar to that obtained with unimanual movement responses. In Experiment 2, an inactive response apparatus was placed to the left or right of the active response apparatus to provide a referent. For half of the participants, the active and inactive apparatuses were joysticks, and for half they were response boxes with keys. For both response types, an up-right/down-left advantage was evident when the relative position of the active response apparatus was right but not when it was left. That bimanual keypresses yield similar eccentricity and relative location effects to those for unimanual movements is predicted by the salient features coding perspective but not by the end-state comfort hypothesis.     

Hemispace Asymmetries and Laterality Effects in Arm Positioning

Hemispace asymmetries and laterality effects were examined on an arm positioning reproduction task. Sixteen male subjects were asked to reproduce both abductive and adductive positioning movements with the left or right arm within either the left or the right hemispace. Hemispace was manipulated using a 90 degrees head-rotation paradigm. A left hemispace advantage in positioning accuracy was predicted for both left and right arm movements on the grounds that the perceptual-motor control of positioning movements made in left hemispace is primarily mediated by the right hemisphere which is known to be advantageous for tasks which are spatial in nature (Heilman, Bowers, & Watson, 1984). No arm laterality effects were predicted to occur because the proximal musculature involved in the control of arm movements is innervated from both contralateral and ipsilateral cerebral hemispheres (Brinkman & Kuypers, 1973). Results showed that the predicted left hemispace advantage was evident for the right arm on the positioning variability measure alone, whereas it was absent for all other possible conditions on all error measures. Laterality (arm) effects were absent as predicted. The experiment also demonstrated a greater degradation of reproduction performance under the ′crossed" arm-hemispace conditions than under the ′uncrossed" conditions. A plausible explanation for the uncrossed advantage for the task is that under normal conditions, a single hemisphere is primarily responsible for both controlling the contralateral arm and directing attention to the contralateral hemispace, and consequently potential interhemispheric interference is minimized. A clear response bias effect in movement reproduction was also evident as a function of the direction of concurrent arm movement and head rotation. Arm movements made in the same direction as head rotation were systematically undershot in reproduction to a much greater degree than arm movements made in the opposite direction to head rotation.     

The effects of skill demands and object position on the distribution of preferred hand reaches

Performance-based measures of hand preference have been developed as an objective method of examining handedness. Previous research using this method showed that both skill demands and the position of the object in working space affect preferential hand reaching. Specifically, preferred hand reaches predominated in left hemispace, in spite of the biomechanical inefficiency involved in reaching across the body midline. This was mediated by the skill demands, with a higher frequency of preferred hand reaches for tasks requiring more skill. To further examine this issue, we increased the task skill demands. Twenty-two right-handed adults reached for five tools located in an array of five positions in front of them. Participants were required to pick up the tool, pick up and demonstrate how to use it, or pick up and actually use the tool on the materials provided. The results showed that the frequency of right hand reaches was greatest for the tool use condition. This effect was mediated by the position of the object in hemispace, with more right hand reaches occurring for the Use task in left hemispace than the other tasks, in support of our previous work.     

Attenuating the haptic horizontal—vertical curvature illusion

In a number of experiments, blindfolded subjects traced convex curves whose verticals were equal to their horizontal extent at the base. Overestimation of verticals, as compared with horizontals, was found, indicating the presence of a horizontal-vertical illusion with haptic curves, as well as with visible curves. Experiment 1 showed that the illusion occurred with stimuli in the frontal plane and with stimuli that were flat on the table surface in vision and touch. In the second experiment, the stimuli were rotated, and differences between vision and touch were revealed, with a stronger illusion in touch. The haptic horizontal-vertical illusion was virtually eliminated when the stimuli were bimanually touched using free exploration at the body midline, but a strong illusion was obtained when curves were felt with two index fingers or with a single hand at the midline. Bimanual exploration eliminated the illusion for smaller 2.5- through 10.2-cm stimuli, but a weakened illusion remained for the largest 12.7-cm patterns. The illusion was present when the stimuli were bimanually explored in the left and right hemispace. Thus, the benefits of bimanual exploration derived from the use of the two hands at the body midline combined with free exploration, rather than from bimanual free exploration per se. The results indicate the importance of haptic exploration at the body midline, where the body can serve as a familiar reference metric for size judgments. Alternative interpretations of the results are discussed, including the impact of movement-based heuristics as a causal factor for the illusion. It was suggested that tracing the curve’s peak served to bisect the curve in haptics, because of the change in direction.     

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.     

Must egocentric and environmental frames of reference be aligned to produce spatial S-R compatibility effects?

Four experiments were conducted to determine the effects of misaligning egocentric and environmental frames of reference on spatial S-R compatibility effects. In Experiments 1 and 3, subjects looked at two lights that were aligned horizontally, one each on either side of the body midline. They held their head upright or tilted 90 degrees to the left or right. In the upright condition the hands were uncrossed and rested opposite the lights (frames of reference aligned), whereas in the head tilt condition the hands were either crossed or uncrossed but positioned perpendicular to the lights (frames of reference not aligned). Manual choice reaction times to the lights produced spatial S-R compatibility effects that were as large when the frames of reference were aligned as when they were not. In Experiments 2 and 4, which also used upright and tilted conditions, we found generally similar results when the lights were displayed vertically and the hands disposed horizontally. The results indicate that under conditions of head rotation and with stimulus and response arrays perpendicular to each other, spatial S-R compatibility effects still occur. By taking into account both frames of reference, the subject classifies the stimuli as left or right whether they are horizontally or vertically disposed and maps them onto the responding hand, thereby producing the observed compatibility effects.     

Influences of hand posture and hand position on compatibility effects for up-down stimuli mapped to left-right responses: evidence for a hand referent hypothesis

Unimanual left-right responses to up-down stimuli show a stimulus-response compatibility (SRC) effect for which the preferred mapping varies as a function of response eccentricity. Responses made in the right hemispace and, to a lesser extent, at a midline position, are faster with the up-right/down-left mapping than with the up-left/down-right mapping, but responses made in the left hemispace are faster with the up-left/down-right mapping. Also, for responses at the midline position, the preferred mapping switches when the hand is placed in a supine posture instead of the more usual prone posture. The response eccentricity effect can be explained in terms of correspondence of asymmetrically coded stimulus and response features, but it is not obvious whether the hand posture effect can be explained in a similar manner. The present study tested the implications of a hypothesis that the body of the hand provides a frame of reference with respect to which the response switch is coded as left or right. As was predicted by this hand referent hypothesis, Experiment 1 showed that the influence of hand posture (prone and supine) on orthogonal SRC was additive with that of response location. In Experiment 2, the location of the switch relative to the hand was varied by having subjects use either a normal grip in which the switch was held between the thumb and the index finger or a grip in which it was held between the little and the ring fingers. The magnitudes of the mapping preferences varied as a function of the grip and hand posture in a manner consistent with the hand referent hypothesis.