Influence of "prehistory" of sequential movements of the right and the left hand on reproduction: coding of positions, movements and sequence structure

The dependence of errors during reproduction of a sequence of hand movements without visual feedback on the previous right- and left-hand performance ("prehistory") and on positions in space of sequence elements (random or ordered by the explicit rule) was analyzed. It was shown that the preceding information about the ordered positions of the sequence elements was used during right-hand movements, whereas left-hand movements were performed with involvement of the information about the random sequence. The data testify to a central mechanism of the analysis of spatial structure of sequence elements. This mechanism activates movement coding specific for the left hemisphere (vector coding) in case of an ordered sequence structure and positional coding specific for the right hemisphere in case of a random sequence structure.     

Motivation and motor control: hemispheric specialization for approach motivation reverses with handedness

Background

According to decades of research on affective motivation in the human brain, approach motivational states are supported primarily by the left hemisphere and avoidance states by the right hemisphere. The underlying cause of this specialization, however, has remained unknown. Here we conducted a first test of the Sword and Shield Hypothesis (SSH), according to which the hemispheric laterality of affective motivation depends on the laterality of motor control for the dominant hand (i.e., the “sword hand," used preferentially to perform approach actions) and the nondominant hand (i.e., the “shield hand," used preferentially to perform avoidance actions).

Methodology/Principal Findings

To determine whether the laterality of approach motivation varies with handedness, we measured alpha-band power (an inverse index of neural activity) in right- and left-handers during resting-state electroencephalography and analyzed hemispheric alpha-power asymmetries as a function of the participants'' trait approach motivational tendencies. Stronger approach motivation was associated with more left-hemisphere activity in right-handers, but with more right-hemisphere activity in left-handers.

Conclusions

The hemispheric correlates of approach motivation reversed between right- and left-handers, consistent with the way they typically use their dominant and nondominant hands to perform approach and avoidance actions. In both right- and left-handers, approach motivation was lateralized to the same hemisphere that controls the dominant hand. This covariation between neural systems for action and emotion provides initial support for the SSH.     

Effects of handedness on visual sensitivity in perihand space

Recent studies have shown that changes in visual processing in perihand space are limited to the area around the right hand, at least in right-handers. One explanation for these findings is that perception is altered at locations where action is more likely to occur. To test this notion, we asked both right- and left-handers to perform an unspeeded visual discrimination task under four hand-position configurations: Left hand, right hand, both hands, or no hands near the display. Compared to the no-hands (control) condition, visual sensitivity (d') was higher in the dominant-hand condition for right-handers and higher in the dominant- as well as the non-dominant hand condition for left-handers. When both hands were near the display, sensitivity was similar to that in the dominant-hand condition for right-handers and to that in the non-dominant hand condition for left-handers. This shows that performance differed between the two handedness groups when their non-dominant hand was near the display (both alone and accompanied by their dominant hand). Thus, the pattern for left-handers did not correspond to a mirror image of the pattern for right-handers. In line with studies on bimanual action control, visual processing in perihand space seems to be determined by the different ways in which left- and right-handers use their hands.     

Evaluation of hand asymmetry in relation to hand preference

We evaluated the asymmetric hand measurements in right- and left-handed individuals. 343 men and 290 women aged 18-42 years (22.11 +/- 2.07) participated in the study. There were no statistically significant differences when right-left differences in hand length, third finger length, palmar length, and the digit index value were evaluated according to hand preference and sex. Statistically significant differences were found for right-left differences in hand width, hand-shape index, and the palmar length/width according to hand preference. The strong left-handers, weak left-handers, and ambidextrous individuals in the study group all exhibited asymmetry favoring the left and were considered together. Similarly, the strong and weak right-handers exhibited asymmetry favoring the right hand and were considered together. The difference between these two groups was significant. When the data were evaluated according to sex, significant differences were found between the subgroups. In particular, right-left differences in the hand-shape index and palmar length/width values of the strong left-handers, weak left-handers, and ambidextrous individuals were found to be statistically significant according to sex; in contrast, the strong and weak right-handers showed no significant differences according to sex. These results suggest a relation of hand asymmetry to hand preference in a Turkish population.     

Bodily Illusions in Young Children: Developmental Change in Visual and Proprioceptive Contributions to Perceived Hand Position

We examined the visual capture of perceived hand position in forty-five 5- to 7-year-olds and in fifteen young adults, using a mirror illusion task. In this task, participants see their left hand on both the left and right (by virtue of a mirror placed at the midline facing the left arm, and obscuring the right). The accuracy of participants’ reaching was measured when proprioceptive and visual cues to the location of the right arm were put into conflict (by placing the arms at different distances from the mirror), and also when only proprioceptive information was available (i.e., when the mirror was covered). Children in all age-groups (and adults) made reaching errors in the mirror condition in accordance with the visually-specified illusory starting position of their hand indicating a visual capture of perceived hand position. Data analysis indicated that visual capture increased substantially up until 6 years of age. These findings are interpreted with respect to the development of the visual guidance of action in early childhood.     

Reaction time in bimanual simultaneous motions

Electromyographically determined reaction times (EMG-RTs) of the finger flexor and extensor of both forearms were measured for four different motions: inward (task 1), flexion of both wrists; outward (task 2), extension of both wrists; to the left (task 3), extension of the left wrist and flexion of the right; and to the right (task 4), flexion of the left and extension of the right. The EMG-RTs were shorter and synchronization errors in terms of left to right differences of EMG-RTs were smaller in tasks 1 and 2 than in tasks 3 and 4. Comparing the flexors and the extensors, the extent of prolongation of EMG-RTs in tasks 3 and 4 differed significantly on the left side, being larger in the flexor than in the extensor, but there was no difference in the extent of prolongation between the flexor and the extensor on the right. It was suggested that the timing of initiation of movements in simultaneous motions was primarily determined by the pattern of muscle coupling in both limbs, and not by the direction of movements. The steadiness of motor function and of the right hand in right-handed subjects was also discussed in regard to hand preference.     

On the Problem of the Leading Activity in Adolescence

The role of interhemispheric asymmetry in the conscious control of sensorimotor measurements of the Muller-Lyer illusion was investigated. We used this perceptual illusion as a phenomenon of consciousness to determine how error value depends on the contribution of the left and right hemisphere to the control of decision making. We compared the precision of the left- and right-hand movements on the touch screen during the measurement of segment length. It was found that the left and right hands controlled by two hemispheres in different ways made different errors during these measurements. The constant bias of right-hand movements is greater than that of the left hand. The conclusion is that the hemisphere primarily involved in movement control determines the dominant system of representation (metric vs. categorical) and the control type in decision making (automatic vs. conscious). The increase or decrease in accuracy of measurement of the illusory object depends on the increase or decrease of conscious control.     

Interhemispheric Inhibition during Mental Actions of Different Complexity

Several investigations suggest that actual and mental actions trigger similar neural substrates. Yet, neurophysiological evidences on the nature of interhemispheric interactions during mental movements are still meagre. Here, we asked whether the content of mental images, investigated by task complexity, is finely represented in the inhibitory interactions between the two primary motor cortices (M1s). Subjects’ left M1 was stimulated by means of transcranial magnetic stimulation (TMS) while they were performing actual or mental movements of increasing complexity with their right hand and exerting a maximum isometric force with their left thumb and index. Thus, we simultaneously assessed the corticospinal excitability in the right opponent pollicis muscle (OP) and the ipsilateral silent period (iSP) in the left OP during actual and mental movements. Corticospinal excitability in right OP increased during actual and mental movements, but task complexity-dependent changes were only observed during actual movements. Interhemispheric motor inhibition in the left OP was similarly modulated by task complexity in both mental and actual movements. Precisely, the duration and the area of the iSP increased with task complexity in both movement conditions. Our findings suggest that mental and actual movements share similar inhibitory neural circuits between the two homologous primary motor cortex areas.     

Recovery of motor skills after the caudate lesion in rats with the different prefered forelimb: the role of intense retraining

Rats were trained for instrumental reaching (17 mm) of a sunflower seed from a horizontal tube 12 mm in diameter. After training, rats were divided into groups, "right-" and "left-handers", by the forelimb preference. Unilateral electrolytic lesions of the head of the caudate nucleus contralateral to the prefered forelimb were performed. After surgery, animals were retrained to perform the food-retrieval reaction by the same forelimb (reaching reaction by the "intact" forelimb was prohibited by a special bracelet). Both right- and left-handers were divided into groups of rare and intense retraining. Over the course of five months, animals of the groups of rare training were tested once a week, whereas the intense retraining was performed three or four times a week. Rats with right and left forelimb preference were shown to recover the reaching skill with different rates. In general, animals with left-side caudate lesion (right-handers) recovered the skill better both under condition of spontaneous recovery (rare testing) and intense retraining. The results suggest different mechanism of skill recovery after the right- and left-side brain lesion.     

Influence of Action-Effect Associations Acquired by Ideomotor Learning on Imitation

According to the ideomotor theory, actions are represented in terms of their perceptual effects, offering a solution for the correspondence problem of imitation (how to translate the observed action into a corresponding motor output). This effect-based coding of action is assumed to be acquired through action-effect learning. Accordingly, performing an action leads to the integration of the perceptual codes of the action effects with the motor commands that brought them about. While ideomotor theory is invoked to account for imitation, the influence of action-effect learning on imitative behavior remains unexplored. In two experiments, imitative performance was measured in a reaction time task following a phase of action-effect acquisition. During action-effect acquisition, participants freely executed a finger movement (index or little finger lifting), and then observed a similar (compatible learning) or a different (incompatible learning) movement. In Experiment 1, finger movements of left and right hands were presented as action-effects during acquisition. In Experiment 2, only right-hand finger movements were presented during action-effect acquisition and in the imitation task the observed hands were oriented orthogonally to participants’ hands in order to avoid spatial congruency effects. Experiments 1 and 2 showed that imitative performance was improved after compatible learning, compared to incompatible learning. In Experiment 2, although action-effect learning involved perception of finger movements of right hand only, imitative capabilities of right- and left-hand finger movements were equally affected. These results indicate that an observed movement stimulus processed as the effect of an action can later prime execution of that action, confirming the ideomotor approach to imitation. We further discuss these findings in relation to previous studies of action-effect learning and in the framework of current ideomotor approaches to imitation.     

Abnormal Movement Preparation in Task-Specific Focal Hand Dystonia

Electrophysiological and behavioral studies in primary dystonia suggest abnormalities during movement preparation, but this crucial phase preceding movement onset has not yet been studied specifically with functional magnetic resonance imaging (fMRI). To identify abnormalities in brain activation during movement preparation, we used event-related fMRI to analyze behaviorally unimpaired sequential finger movements in 18 patients with task-specific focal hand dystonia (FHD) and 18 healthy subjects. Patients and controls executed self-initiated or externally cued prelearnt four-digit sequential movements using either right or left hands. In FHD patients, motor performance of the sequential finger task was not associated with task-related dystonic posturing and their activation levels during motor execution were highly comparable with controls. On the other hand reduced activation was observed during movement preparation in the FHD patients in left premotor cortex / precentral gyrus for all conditions, and for self-initiation additionally in supplementary motor area, left mid-insula and anterior putamen, independent of effector side. Findings argue for abnormalities of early stages of motor control in FHD, manifesting during movement preparation. Since deficits map to regions involved in the coding of motor programs, we propose that task-specific dystonia is characterized by abnormalities during recruitment of motor programs: these do not manifest at the behavioral level during simple automated movements, however, errors in motor programs of complex movements established by extensive practice (a core feature of FHD), trigger the inappropriate movement patterns observed in task-specific dystonia.     

Handedness in comparison with the asymmetry of the upper extremities]

In 1990 221 males (88 children, age 3.5-7.25 years and 133 adults, age 20-44 years) were investigated. The sample was divided into two groups based on handedness; 140 individuals were right-handed and 81 left-handed. 10 measurements were taken at the upper limbs. In addition, marked handedness, handclasping and armfolding were determined. The last three traits showed no statistically significant connection. It was determined that measurements of length, breadth and circumference are more often significantly asymmetric in favor of the right side in right-handers than in left-handers. Further, skinfolds of right-handed individuals are larger on the left side. Partially reverse results were found for left-handers as a whole, however, the results are more irregular. A difference in the degree of asymmetry exists between children and adults. Asymmetries of breadth and circumference measurements as well as forearm skinfold are larger in adults. Taking into account the error of measurement, frequencies of R > L, R = L and L > R for right- and left-handed individuals were evaluated using the chi 2-test for significance. Preponderance of measurements on the dominant side could often be secured statistically.     

Prevalence of the three categories of handedness among Malawian school children

A cross sectional study of prevalence of left, right and mixed handedness was made on 512 Malawian school children (240 boys and 272 girls) aged 6-17 years. Handedness was assessed using questionnaire incorporating questions on hand preferred for eight unimanual activities. Interviews with guardians, pupils and teachers were conducted to assess the pressure experienced by children to use right hand for unimanual activities. The overall prevalence of left-, right- and mixed-handers was 3.9%, 90.4% and 5.7%, respectively. Association between handedness and age or gender of children was nonsignificant. 96.2% of guardians and 92.7% of teachers were insistent upon use of right hand for unimanual tasks. Most of non-right handed children indicated that they had experienced pressure to use right hand for unimanual activities but they were still using left hand for one or more manual tasks. 28 pupils indicated that they stopped using left hand under the pressure from guardians and teachers.     

Adjustment errors of feedback guided human forearm movements elicited by random position signal sequences

The effects of visual and auditory relevant feedback on human forearm movements elicited by random position signal sequences were examined. We observed the adjustment error sequences when the arm moved briskly and accurately to the target position. As the results, the mean adjustment errors for each speed (clock interval) of random position signal sequence are different in the left-right target positions, and also different in the flexion-extension movements. In faster speed (1s), the adjustment errors are approximately flat as compared with those in other speeds, on the other hand, in moderate speed (2s) or slower speed (3s), these indicate a upward tendency from left to right positions, and also indicate a upward tendency from flexion to extension movements. However, in the reaction times, the moving times, and the adjusting times, there are no significant differences for the left-right target positions and for the flexion-extension movements.     

Impaired mirror-image imitation in Asperger and high-functioning autistic subjects

Imitation is crucial for proper development of social and communicative skills. Here, we argue that, based on an error analysis of a behavioral imitation task, adult Asperger and high-functioning autistic subjects suffer from an intriguing deficit of imitation: they lack the natural preference for imitation in a mirror-image fashion. The imitation task consisted of a simple movement sequence of putting a pen with the left or right hand into a green or a blue cup using one of two possible grips. The subjects were asked to imitate the experimenter's hand movements either using the crossed hand (e.g., the subject's right hand corresponding to the experimenter's right hand) for imitation or to imitate as if looking in a mirror (e.g., the subject's left hand corresponding to the experimenter's right hand). When people normally view other persons face-to-face, they prefer to imitate as in a mirror, and observation of mirror-image-like movements speeds up performance in nonimitative tasks. However, our autistic subjects, defective in social cognition, did not profit from mirror-image movements of others. These results provide a new insight into the difficulties that autistic subjects face in viewing and understanding actions of others.     

Cerebral haemodynamics during motor imagery of self-feeding with chopsticks: differences between dominant and non-dominant hand

Abstract

Purpose: Motor imagery is defined as a dynamic state during which a subject mentally simulates a given action without overt movements. Our aim was to use near-infrared spectroscopy to investigate differences in cerebral haemodynamics during motor imagery of self-feeding with chopsticks using the dominant or non-dominant hand.

Materials and methods: Twenty healthy right-handed people participated in this study. The motor imagery task involved eating sliced cucumber pickles using chopsticks with the dominant (right) or non-dominant (left) hand. Activation of regions of interest (pre-supplementary motor area, supplementary motor area, pre-motor area, pre-frontal cortex, and sensorimotor cortex was assessed.

Results: Motor imagery vividness of the dominant hand tended to be significantly higher than that of the non-dominant hand. The time of peak oxygenated haemoglobin was significantly earlier in the right pre-frontal cortex than in the supplementary motor area and left pre-motor area. Haemodynamic correlations were detected in more regions of interest during dominant-hand motor imagery than during non-dominant-hand motor imagery.

Conclusions: Haemodynamics might be affected by differences in motor imagery vividness caused by variations in motor manipulation.     

Mental representations of movements. Brain potentials associated with imagination of hand movements

The present study was designed in order to contribute towards the understanding of the physiology of motor imagery. DC potentials were recorded when subjects either imagined or executed a sequence of unilateral or bilateral hand movements. The sequence consisted of hand movements in 4 directions, forwards, backwards, to the right and to the left, and varied from trial to trial. The sequence had been cued by visual targets on a computer screen and had to be memorized before the trial was initiated. Changes of DC potentials between task execution and imagination were localized in central recordings (C3, Cz, C4) with larger amplitudes when executing the task than when imagining to do so. Stimulation of peripheral receptors associated with task execution or a different level of activation of the cortico-motoneural system could account for this finding. The main result of the present study was that with unilateral performance, the side of the performing hand (right, left) had localized effects in recordings over the sensorimotor hand area (C3, C4) which were qualitatively the same with imagination and execution and quantitatively similar (i.e., without significant difference). Performance of the right hand augmented negative DC potentials in C3, performance of the left hand augmented amplitudes in C4. This result is consistent with the assumption that the primary motor cortex is active with motor imagery. Finally, the question has been addressed whether motor imagery may involve the left hemisphere to a larger extent than the execution of the movement. It is shown that a particular contribution of the left hemisphere associated with motor imagery may only show up under strictly controlled conditions.     

Selective interference for the right hemisphere in a task of visual-motor exploration

Simple reaction times to lateralized visual stimuli were studied in normal subjects while they were carrying out a concomitant task. The concomitant task consisted in the exploration of a visual maze presented in the middle of a screen. Regardless of the hand used, the concomitant task produced a specific lengthening of the responses to stimuli located in the left visual field. It is concluded that the right hemisphere plays a major role in the organization of ocular movements during active exploration of visual environment.     

Hand preferences of captive chimpanzees (Pan troglodytes) in simple reaching for food

We examined chimpanzee hand preference in simple reaching for food, with special reference to manipulative patterns and the developmental shift. We observed 80 captive chimpanzees, ranging from 1 to 25 years old. We also studied the manipulative patterns (grip- types) of 70 individuals as they reached for raisins scattered randomly on the floor. We employed LQ score as a measure of hand preference and designated the subjects right- handers (or left- handers) if they used their right hands (left hands) above chance level. Although the numbers of right- handers and left- handers are almost equal, the distribution of the strength is not symmetrical in both groups. Strong preference was exhibited by more left- handers than right- handers. Subjects > 9 years old exhibited greater hand preference, whereas subjects < 9 years old were ambidextrous. We classified manipulative patterns for reaching into five basic grip- types and analyzed them vis- à- vis age. There is no significant correlation between preferred hand and manipulative patterns. However, adult subjects tended to use an index- and - middle- finger grip with the left hand and to use imprecise grips with the right hand more often than other patterns regardless which hand they preferred. These data demonstrate a developmental shift in hand preference and manipulative patterns and also reveal functional asymmetries between the right and the left hand in Pan troglodytes.     

Hand preferences of captive chimpanzees (Pan troglodytes) in simple reaching for food

We examined chimpanzee hand preference in simple reaching for food, with special reference to manipulative patterns and the developmental shift. We observed 80 captive chimpanzees, ranging from 1 to 25 years old. We also studied the manipulative patterns (grip- types) of 70 individuals as they reached for raisins scattered randomly on the floor. We employed LQ score as a measure of hand preference and designated the subjects right- handers (or left- handers) if they used their right hands (left hands) above chance level. Although the numbers of right- handers and left- handers are almost equal, the distribution of the strength is not symmetrical in both groups. Strong preference was exhibited by more left- handers than right- handers. Subjects > 9 years old exhibited greater hand preference, whereas subjects < 9 years old were ambidextrous. We classified manipulative patterns for reaching into five basic grip- types and analyzed them vis- à- vis age. There is no significant correlation between preferred hand and manipulative patterns. However, adult subjects tended to use an index- and - middle- finger grip with the left hand and to use imprecise grips with the right hand more often than other patterns regardless which hand they preferred. These data demonstrate a developmental shift in hand preference and manipulative patterns and also reveal functional asymmetries between the right and the left hand in Pan troglodytes.