The Specificity of Motor Skill and Manual Asymmetry: A Review of the Evidence and Its Implications

Results of investigations on gaining control of limb movements are reviewed, and their contribution to understanding the development of manual asymmetries is discussed in relation to the discrimination and programming of appropriate neuromuscular resources. An examination of the relevant evidence on number and types of manual asymmetries recorded provides strong grounds for concluding that where asymmetries occur, they simply represent a further example of the well-documented activity-specific nature of motor skills and of the extremely lengthy periods of learning or experience needed for their acquisition and perfection. This specificity of motor skill and manual asymmetry also readily accounts for most of the discrepancy usually reported between assessments of hand preference and performance differences between hands, because these alternative measures of handedness have rarely employed the same range or variety of tasks.     

Speech fluency and hand performance on a sequential tapping task in left- and right-handed stutterers and nonstutterers

The purpose of this study was to examine hemispheric functioning in right- and left-handed stutterers and nonstutterers using tasks that require concurrent speech and unimanual sequential tapping activity. Twenty adult stutterers (15 right-handers and 5 left-handers) and 20 adult nonstutterers (15 right-handers and 5 left-handers) performed four experimental tasks: tapping, tapping-spontaneous speech, tapping-reading, and tapping-singing. Subjects performed each task for 120 sec (8 trials of 15 sec per trial), alternating tapping hands after each trial.Results indicated that subjects' speech and manual performance were hindered by the concurrent motor activities and that stutterers demonstrated more interference than the nonstutterers. In addition, the tapping-spontaneous speech task resulted in the greatest interfering condition. Results suggest that both intrahemispheric competition and interhemispheric integration processes are involved in the regulation of speech and motor activities. Moreover, the possibility exists that a key difference between stutterers and nonstutterers may be linked to the degree to which these activities are synchronized.     

Menstrual cycle-related changes of functional cerebral asymmetries in fine motor coordination

Fluctuating sex hormone levels during the menstrual cycle have been shown to affect functional cerebral asymmetries in cognitive domains. These effects seem to result from the neuromodulatory properties of sex hormones and their metabolites on interhemispheric processing. The present study was carried out to investigate whether functional cerebral asymmetries in fine motor coordination as reflected by manual asymmetries are also susceptible to natural sex hormonal variations during the menstrual cycle. Sixteen right-handed women with a regular menstrual cycle performed a finger tapping paradigm consisting of two conditions (simple, sequential) during the low hormone menstrual phase and the high estrogen and progesterone luteal phase. To validate the luteal phase, saliva levels of free progesterone (P) were analysed using chemiluminescence assays. As expected, normally cycling women showed a substantial decrease in manual asymmetries in a more demanding sequential tapping condition involving four fingers compared with simple (repetitive) finger tapping. This reduction in the degree of dominant (right) hand manual asymmetries was evident during the luteal phase. During the menstrual phase, however, manual asymmetries were even reversed in direction, indicating a slight advantage in favour of the non-dominant (left) hand. These findings suggest that functional cerebral asymmetries in fine motor coordination are affected by sex hormonal changes during the menstrual cycle, probably via hormonal modulations of interhemispheric interaction.     

Unilateral agraphia after section of the posterior half of the truncus of the corpus callosum

Children ranging in age from 3 to 12 tapped on a morse key with their index finger as fast as possible. After a control test, they concurrently tapped and recited a nursery rhyme, recited animal names, and memorized shapes. Previous results suggested that more interference in a motor task results from attempting at the same time to perform another task controlled by the same, than by a different, hemisphere. Right-handed tapping and talking both used left hemisphere space; left-handed tapping and talking were controlled by different hemispheres. Relative to silent tapping concurrent talking caused a greater drop in right-hand tapping rate than in left-hand tapping rate in both the rhyme and animal conditions. Interference by shape memorizing was equal across hands. The differential effect of speaking on right-hand preference indicates the left lateralization of speech output control. The size of this effect did not vary with increasing grade level. The findings support the view that speech output control is fully lateralized at least by age 3 years.     

The influence of dominant versus non-dominant hand on event and emergent motor timing

It has been hypothesized that timing in tapping utilizes event timing; a clock-like process, whereas timing in circle drawing is emergent. Three experiments examined timing in tapping and circle drawing by the dominant and non-dominant hand. Participants were right-hand dominant college aged males and females. The relationship between variance and the square of the timed interval (the Weber fraction), thought to capture clock-like timekeeping processes, was compared. Furthermore, timing variance was decomposed into a clock and a motor component. The slopes for timing were different for dominant hand tapping and circle drawing, but equal for non-dominant and dominant hand tapping. Negative lag one covariance, consistent with motor implementation variability, was found for non-dominant but not for dominant hand circle drawing (Experiment 1). Practice did not influence this relation (Experiment 2). A significant correlation for clock variability was found between non-dominant hand circle drawing and tapping (Experiment 3). Collectively, these findings indicate that event timing is shareable across hands while emergent timing is specific to an effector. Emergent timing does not appear to be obligatory for the non-dominant hand in circle drawing. We suggest that the use of emergent timing might depend upon the extensive practice experienced by a person's dominant hand.     

Hemispheric interference: A dual-task investigation of youngsters who stutter

Hemispheric interference was studied in this investigation by requiring 24 right-handed grade-school children who stutter to tap a button, as rapidly as possible, in five randomized conditions. Hand order was counterbalanced in the single-task silent control condition and in the four increasingly demanding dual-task conditions. In three of the experimental conditions tapping and speech occured concurrently. In the fourth dual-task condition the subjects were to vocalize the sound of a siren (in an irregularly patterned way) as they tapped the button. Analysis of the tapping rates in this two-by-five factorial study revealed statistically significant differences among hands and conditions. The tapping rate was faster with the right than with the left hand and the rate decreased as the demands of the dual-task conditions increased. Interaction was not evidenced; the interference pattern was similar in both hands across the conditions. Since interference was significantly present in both the speech and vocalic conditions, the data are not consistent with a laterality model. This finding and the increase in interference with increased attentional demand, however, are consistent with a capacity framework.     

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

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

Bimanual coordination dynamics in poststroke hemiparetics

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

The contribution of tactile reafference to temporal regularity during bimanual finger tapping

In a repetitive tapping task, the within-hand variability of intertap intervals is reduced when participants tap with both hands, as opposed to single-handed tapping. This bimanual advantage can be attributed to timer variance (according to the Wing-Kristofferson model). Separate timers have been proposed for each hand whose outputs are then averaged (Helmuth & Ivry, 1996, Journal of Experimental Psychology: Human Perception and Performance, 22, 278-293). Alternatively, timing might be based on sensory reafference and the bimanual advantage due to the enhancement of sensory reafferences. This alternative hypothesis was tested in three experiments. In the first experiment, we replicated the bimanual advantage in tapping with two fingers of the same hand compared with single finger tapping. In the second experiment, we demonstrated that the bimanual advantage decreased when tactile reafferences from left-hand taps were omitted (by contact-free tapping). In the third experiment, participants tapped bimanually with the index fingers of both hands firmly mechanically coupled. The bimanual advantage was replicated for this condition. Results are consistent with the assumption that the bimanual advantage is due to the sensory reafferences of the second hand. We suggest that our results are best explained by a reformulation of the Wing-Kristofferson model, in which the timer provides action goals in terms of sensory reafferences.     

Roles of Visual Information for Control of Reaching Movements in Children

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

Eye movements and performance during bilateral tracing tasks

To examine the role of current visual monitoring in the between-hand differences in skilled movements, eye movements and errors during bilateral tracing tasks were analyzed in 10 subjects. When subjects traced a horizontal course abductively with both hands, the subject's gaze followed the movement of the right hand, and more errors were observed on the left hand. When subjects traced a vertical course, where fine motor control for alteration of movement direction was necessary, more errors were shown on the left hand despite the alternate visual scan of the two hands. The results were interpreted as showing that the between-hand differences in skilled movements are primarily due to the left hand's poor ability in motor output, and that the differential efficiency in the use of visual monitoring becomes an important factor in the between-hand differences when symmetrical movements of both hands with a low degree of difficulty are required.     

Hemispheric asymmetries in attentional control: implications for hand preference in sensorimotor tasks

Liepmann (1908) proposed that handedness reflects the greater capacity of one hemisphere to learn the execution of skilled movements. Although asymmetries in motor control are an important basis for hand asymmetries, recent studies have suggested that handedness may be determined by multiple factors. In the present study, we examine how attentional asymmetries may contribute to hand preferences. Right-handed subjects participated in a reaction time task in which they were given preliminary information about where a target stimulus would occur (selective attention) or which hand to use for responding (selective intention). Our findings indicate that these processes influence each other reciprocally and favor a state of optimal attentional and intentional preparation of the right hand. We suggest that these hemispheric asymmetries in attentional control contribute to hand preferences in certain sensorimotor tasks.     

Laterality differences for speed but not for control in sequential finger tapping

Multiple or sequential finger tapping is preferential to the dominant right hand with respect to speed. However, in more complex movement, variables other than speed become important. The present investigation uses a sequential finger-tapping task which permits assessment of between-hands differences with respect to rate and control of movement, with and without vision. 36 right-handed normal adults rapidly tapped their fingers in sequential order on a block (2.54 cm. sq.), trying not to move the block. Analyses of variance (mode x hand) performed for taps and shift of the block show the right hand to be faster than the left hand with and without vision, adding further to the notion that the left hemisphere predominates in increases in rapid movement and in sequencing aspects of motor activity. However, while both hands were steadier with vision than without, there were no between-hand differences with regard to control, suggesting equivalency of cerebral function for factors of manual sequencing other than speed.     

Relationship between extremities in motor performance

The purpose of the study was to examine the relationship between the extremities in different aspects of motor performance, including simple reaction time, choice reaction time, speed of movement, tapping speed, and coordination, i.e., speed of movement/accuracy. 186 healthy randomly selected right-handed subjects (93 men, 93 women; aged 21-70 years) were measured on a test battery of five tests for both hands and feet. Correlations between the extremities in motor performance were highest between hands (r=.71-.80, p<.001) or between feet (r=.57-.88, p<.001) depending on the complexity of response for all subjects. These relations remained at almost the same magnitude even when age was eliminated. Correlations between the upper and lower extremities were lower (r = .40-.62). Correlations between extremities were lower for one age group (21-30 years) than for all subjects (21-70 years). These results showed that the motor performance for the hands is not a very good indicator of the motor performance of the lower extremities, especially in a homogeneous age group, and it seems that the lower extremities should be studied with specific motor performance tests.     

Perceptual and motor factors in the imitation of simple temporal patterns

Summary This study investigated the relative importance of perceptual and motor factors in the imitation of simple temporal patterns. Previous research in which subjects tap out interval sequences using one finger has suggested that perceptual factors play an important role in response timing. Studies of bimanual tapping, in contrast, stress the importance of motor interactions between the two hands. In this experiment we compared the ability of subjects to tap out two-interval sequences using one finger, two fingers on one hand, and two fingers on opposite hands. The results showed almost identical performance under the three response conditions. It is suggested that the perceptual relations between intervals in a pattern were the main determinant of performance in this experiment.     

Handedness and Hemispheric Asymmetry in the Control Of Movements

This study investigated the influence of hemispheric specialization of function on the motor performance of the hands. Right-handed (n=17), ambidextral (n=21), and ambisinistral (n=12) subjects performed Fitts' (1954) reciprocal tapping task under two conditions with each hand. Conditions had the same index of difficulty but differed in movement precision. The left hand of righthanders was superior in the condition requiring the greatest amount of preprogramming. Conversely, the right hand was not superior in the condition having the greatest demand for feedback control. For ambidextrals, left-hand superiority in the relatively preprogrammed condition was also revealed. Ambisinistrals showed no significant difference between conditions with either hand. The results partially support the hypothesis that the motor performance of the hands mirrors the dominant processing mode of their contralateral hemisphere. Failure to find supportive evidence in ambisinistrals is consistent with the contention that they lack hemispheric specialization of function.     

Temporal Regularity of Tapping by the Left and Right Hands in Timed and Untimed Finger Tapping

The temporal characteristics of repetitive finger tapping by the left and right hands were examined in two experiments. In the first experiment, interresponse intervals (IRIs) were recorded while right-handed male subjects tapped in synchrony with an auditory timing pulse (the synchronization phase) and then attempted to maintain the same tapping rate without the timing pulses (the continuation phase). The left and right hands performed separately, at four different rates (interpulse intervals of 250, 500, 750, and 1500 ms). There was no asymmetry of the asynchronies of the timing pulses and the associated responses in the synchronization phase or of the IRIs in either phase, but there was an asymmetry in the temporal dispersion of the responses in both phases. In the second experiment, right-handed males tapped separately with each hand at three different speeds: as quickly as possible, at a fast but steady rate, and at a slow rhythmical rate. The speed asymmetry present when tapping as quickly as possible (with the preferred hand tapping more quickly) was reduced when tapping at the fast steady rate and was absent when tapping at the slow rhythmical rate. The temporal dispersion of the IRIs produced by the nonpreferred hand was greater than the temporal dispersion of those produced by the preferred hand in all speed conditions. These results show smaller temporal dispersion of tapping by the preferred hand in right-handed males under different conditions, including submaximal speeds at which both hands respond at the same rate. This suggests that the motor system controlling the preferred hand in right-handers has more precise timing of response output than that controlling the nonpreferred hand.     

Handedness and hemispheric asymmetry in the control of movements

This study investigated the influence of hemispheric specialization of function on the motor performance of the hands. Right-handed (n=7), ambidextral (n=21), and ambisinistral (n=12) subjects performed Fitts' (1954) reciprocal tapping task under two conditions with each hand. Conditions had the same index of difficulty but differed in movement precision. The left hand of right-handers was superior in the condition requiring the greatest amount of preprogramming. Conversely, the right hand was not superior in the condition having the greatest demand for feedback control. For ambidextrals, left-hand superiority in the relatively preprogrammed condition was also revealed. Ambisinistrals showed no significant difference between conditions with either hand. The results partially support the hypothesis that the motor performance of the hands mirrors the dominant processing mode of their contralateral hemisphere. Failure to find supportive evidence in ambisinistrals is consistent with the contention that they lack hemispheric specialization of function.     

Handedness and Reach-to-Place Kinematics in Adults: Left-Handers Are Not Reversed Right-Handers

The primary goal of this study was to examine the relations between limb control and handedness in adults. Participants were categorized as left or right handed for analyses using the Edinburgh Handedness Inventory. Three-dimensional recordings were made of each arm on two reach-to-place tasks: adults reached to a ball and placed it into the opening of a toy (fitting task), or reached to a Cheerio inside a cup, which they placed on a designated mark after each trial (cup task). We hypothesized that limb control and handedness were related, and we predicted that we would observe side differences favoring the dominant limb based on the dynamic dominance hypothesis of motor lateralization. Specifically, we predicted that the dominant limb would be straighter and smoother on both tasks compared with the nondominant limb (i.e., right arm in right-handers and left arm in left-handers). Our results only partially supported these predictions for right-handers, but not for left-handers. When differences between hands were observed, the right hand was favored regardless of handedness group. Our findings suggest that left-handers are not reversed right-handers when compared on interlimb kinematics for reach-to-place tasks, and reaffirm that task selection is critical when evaluating manual asymmetries.     

Development of sensorimotor synchronization abilities: Motor and cognitive components

The aim of the present study was to examine both the development of sensorimotor synchronization in children in the age range from 5 to 8 years and the involvement of motor and cognitive capacities. Children performed a spontaneous motor tempo task and a synchronization–continuation task using an external auditory stimulus presented at three different inter-stimulus intervals: 500, 700, and 900 ms. Their motor and cognitive abilities (short-term memory, working memory, and attention) were also assessed with various neuropsychological tests. The results showed some developmental changes in synchronization capacities, with the regularity of tapping and the ability to slow down the tapping rate improving with age. The age-related differences in tapping were nevertheless greater in the continuation phase than in the synchronization phase. In addition, the development of motor capacities explained the age-related changes in performance for the synchronization phase and the continuation phase, although working memory capacities were also involved in the interindividual differences in performance in the continuation phase. The continuation phase is thus more cognitively demanding than the synchronization phase. Consequently, the improvement in sensorimotor synchronization during childhood is related to motor development in the case of synchronization but also to cognitive development with regard to the reproduction and maintenance of the rhythm in memory.