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.     

Preference for Orientations Commonly Viewed for One’s Own Hand in the Anterior Intraparietal Cortex

Brain regions in the intraparietal and the premotor cortices selectively process visual and multisensory events near the hands (peri-hand space). Visual information from the hand itself modulates this processing potentially because it is used to estimate the location of one’s own body and the surrounding space. In humans specific occipitotemporal areas process visual information of specific body parts such as hands. Here we used an fMRI block-design to investigate if anterior intraparietal and ventral premotor ‘peri-hand areas’ exhibit selective responses to viewing images of hands and viewing specific hand orientations. Furthermore, we investigated if the occipitotemporal ‘hand area’ is sensitive to viewed hand orientation. Our findings demonstrate increased BOLD responses in the left anterior intraparietal area when participants viewed hands and feet as compared to faces and objects. Anterior intraparietal and also occipitotemporal areas in the left hemisphere exhibited response preferences for viewing right hands with orientations commonly viewed for one’s own hand as compared to uncommon own hand orientations. Our results indicate that both anterior intraparietal and occipitotemporal areas encode visual limb-specific shape and orientation information.     

Linear Blood Velocity in Arteries of the Brain Hemispheres in Left-Handers and Right-Handers during Hypoxia

Transcranial Doppler ultrasonography was used to measure the linear blood velocity (LBV) in the middle cerebral artery (MCA) of the right and left hemispheres in right-handers and left-handers who were breathing a gas mixture with 10% oxygen through a mask at normal barometric pressure. No interhemispheric differences were found in left-handers before hypoxia. The maximum systolic velocity was higher in the MCA of the left hemisphere in right-handers. Left-handers had a higher LBV in the right hemisphere than did right-handers. LBV in the MCA of both hemispheres increased in both left-handers and right-handers as a result of hypoxia. Aggravation of hypoxia caused a decrease in LBV in the left hemisphere in right-handers.     

Memorization of sequences of right and left hand movements in right- and left-handers: Vector coding

Using studies of the right and left hemisphere’s specialization for positional and vector coding, we analyzed the errors made by right- and left-handers while reproducing sequences of right and left hand movements in a task that activates vector coding by changing the order of movements in memorized sequences. The task was performed first with one hand (starting) and then with the other (continuing). Both right- and lefthanders were found to use information about previous movements of the starting hand only when the dominant hand was starting. After changing the hand, right-handers used information about previous movements of the continuing hand, while left-handers did not. The results were compared with data from earlier experiments wherein positional coding was activated. The comparison showed that vector coding was predominantly involved in memorizing sequences of movements made by the dominant hand, while positional coding was used in the case of the opposite hand in both right- and left-handers. Patterns of errors after changing the hand differed between right- and left-handers, and the conclusion was made that skills are transferred in different ways in right- and left-handers, depending on the type of coding.     

Good and Bad in the Hands of Politicians: Spontaneous Gestures during Positive and Negative Speech

Background

According to the body-specificity hypothesis, people with different bodily characteristics should form correspondingly different mental representations, even in highly abstract conceptual domains. In a previous test of this proposal, right- and left-handers were found to associate positive ideas like intelligence, attractiveness, and honesty with their dominant side and negative ideas with their non-dominant side. The goal of the present study was to determine whether ‘body-specific’ associations of space and valence can be observed beyond the laboratory in spontaneous behavior, and whether these implicit associations have visible consequences.

Methodology and Principal Findings

We analyzed speech and gesture (3012 spoken clauses, 1747 gestures) from the final debates of the 2004 and 2008 US presidential elections, which involved two right-handers (Kerry, Bush) and two left-handers (Obama, McCain). Blind, independent coding of speech and gesture allowed objective hypothesis testing. Right- and left-handed candidates showed contrasting associations between gesture and speech. In both of the left-handed candidates, left-hand gestures were associated more strongly with positive-valence clauses and right-hand gestures with negative-valence clauses; the opposite pattern was found in both right-handed candidates.

Conclusions

Speakers associate positive messages more strongly with dominant hand gestures and negative messages with non-dominant hand gestures, revealing a hidden link between action and emotion. This pattern cannot be explained by conventions in language or culture, which associate ‘good’ with ‘right’ but not with ‘left’; rather, results support and extend the body-specificity hypothesis. Furthermore, results suggest that the hand speakers use to gesture may have unexpected (and probably unintended) communicative value, providing the listener with a subtle index of how the speaker feels about the content of the co-occurring speech.     

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.     

Handedness is a determining factor in lateralized olfactory discrimination

The study aimed to re-investigate differences in olfactory thresholds and odor discrimination between the left and right sides in relation to the handedness of healthy subjects. Twenty left- and 20 right-handers participated; all were in excellent health with no indication of any major nasal or health problems, and all were non-smokers. The two groups were comparable in terms of sex and age (left-handers: 11 women, 9 men, median age 25 years; right-handers: 9 women, 11 men, median age 26 years). Odor thresholds did not differ in relation to handedness. However, in the odor discrimination task the left-handers performed significantly better at the left side compared with the right nostril; this pattern was reversed in the right-handers. The data indicate that, similar to other sensory systems, higher olfactory functions exhibit a certain degree of lateralization.      

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.     

Actigraphic investigations on the activity-rest behavior of right- and left-handed students

The aim of this study was to explore differences between left-and right-handed subjects in sleep duration. Sleep and activity patterns were continuously registered for 12 days using actometers on 20 left-handed and 20 right-handed medical students in Berlin. Handedness was determined by a modified version of the Edinburgh handedness inventory. Each participant wore one actometer on each wrist. Actiwatch^® Sleep Analysis Software (CNT, UK) was used to evaluate the data, and statistical calculations were performed with a non-parametric variance analysis. A significant difference in mean sleep duration between left-handers (7.9 h) and right-handers (7.3 h) was determined (p=0.025 for measurement made on the dominant hand and p=0.013 for ones made on the non-dominant hand). In contrast, the maximal phase of daily activity (acrophase) did not show any difference between the two groups. The difference in sleep duration might be caused by either the greater effort required for left-handers to cope in a right-handed world or by structural brain differences.     

Writing and drawing with both hands as indicators of hemispheric dominance

Brain lateralization is a common term used to describe dominance of one brain hemisphere over another for a specific function. The right hand dominance in writing, controlled by the left hemisphere, is preceded by development of communicative gesticulation and followed by development of speech in the same hemisphere. We assumed that some people are not aware of their own capability of using the other hand for tasks involving fine motor sequential movements. To prove this hypothesis, the participants were asked to perform one trained task (writing) and one less-trained task (drawing) with a dominant and a non-dominant hand. The final sample was comprised of 1189 children from 14 elementary schools and 8 high schools in the Osijek area, of which 685 elementary school children were attending 1st to 4th grade and 504 high school children were attending 3rd and 4th grade. The participants were asked to write two words, draw a specific object (a vase with flowers) and fill out a questionnaire with 10 questions concerning the classification of handedness and cerebral hemisphere dominance. The self-reported cerebral lateralization assessed in the questionnaire was compared with the drawing and the writing performance. The self-reported and objectively measured hand dominance deviated in the cases of the ambidextrous who consider themselves right-handers. Given the fact that the number of ambidextrous persons was greater in elementary schools than in high schools, we concluded how training of the right hand decreases the ability of using both hands equally for either of the tested functions - writing and drawing.     

The practice effect and its difference of the pursuit rotor test with the dominant and non-dominant hands

The purpose of this study was to examine the practice effect and what difference it makes in the pursuit rotor test for the dominant and non-dominant hands in 30 right-handed Japanese male adults aged 18 to 23 years (Age 20.8+/-1.4 yrs). The subjects performed the pursuit rotor test for 1 min in 20 trials with a 1-min interval alternately using the dominant and non-dominant hands. After continuing for 10 trials, a 5-min rest was taken. The measurement order was randomly assigned. Contact time of a steel pen and a target was measured in units of 1/10 sec. The measurements showed a constant increasing tendency at every trial until the 6th trial in both hands. Significant linear regressions were identified, but the increase-rate of the dominant hand was significantly larger. Individual differences showed a decreasing tendency at every trial in the dominant hand, but in the non-dominant hand it increased until the 4th trial then decreased. The relationships between measurements of the 1st and 10th trials in both hands and both hands in the 1st or 10th trial were not high. The performance of the pursuit rotor test improves at every trial in both hands, but the improvement rate decreases after the 7th trial. The improvement rate of the dominant hand is high. The change in individual differences differs in both hands and the relationship between the measurements is not high. It can be judged that the practice effect of the pursuit rotor test differs in the dominant and non-dominant hands.     

Spatial compatibility and position of the effectors

Mean reaction times obtained with crossed hands (right had on the left and left hand on the right) are slower than reaction times obtained with uncrossed hands (right hand on the right and left hand on the left). These results have been explained as a compatibility effect between the responding hand and its spatial position. The goal of the present experiment was to establish whether the position of the hand is encoded by the subjects relative to their body (absolute position) or relative to the other hand (relative position). The subjects performed a discrimination task on two visual stimuli. Stimuli and hands were either on the same side of the body (both on the left or both on the right) or had different absolute position. In all conditions the subjects responded with crossed and uncrossed hands. The results support the hypothesis that relative position is encoded.     

Direct evidence from parietal extinction of enhancement of visual attention near a visible hand

Brain areas exist that appear to be specialized for the coding of visual space surrounding the body (peripersonal space). In marked contrast to neurons in earlier visual areas, cells have been reported in parietal and frontal lobes that effectively respond only when visual stimuli are located in spatial proximity to a particular body part (for example, face, arm or hand) [1-4]. Despite several single-cell studies, the representation of near visual space has scarcely been investigated in humans. Here we focus on the neuropsychological phenomenon of visual extinction following unilateral brain damage. Patients with this disorder may respond well to a single stimulus in either visual field; however, when two stimuli are presented concurrently, the contralesional stimulus is disregarded or poorly identified. Extinction is commonly thought to reflect a pathological bias in selective vision favoring the ipsilesional side under competitive conditions, as a result of the unilateral brain lesion [5-7]. We examined a parietally damaged patient (D.P.) to determine whether visual extinction is modulated by the position of the hands in peripersonal space. We measured the severity of visual extinction in a task which held constant visual and spatial information about stimuli, while varying the distance between hands and stimuli. We found that selection in the affected visual field was remarkably more efficient when visual events were presented in the space near the contralesional finger than far from it. However, the amelioration of extinction dissolved when hands were covered from view, implying that the effect of hand position was not mediated purely through proprioception. These findings illustrate the importance of the spatial relationship between hand position and object location for the internal construction of visual peripersonal space in humans.     

Handedness and socioeconomic status in an urban population in Uzbekistan

The persistence of left-handers in every human population studied to date is an evolutionary puzzle in light of evidence of survival costs associated with left-handedness. Associations between left-handedness and socioeconomic advantages have been observed in Western countries and could provide left-handers fitness benefits through higher survival chances and greater reproductive success. We aimed to explore the generality of this result in another culture. For this purpose, we investigated several socioeconomic status indicators and the number of children alive for 917 men and women in Uzbekistan and compared results for two different measures of handedness: hand preferences for writing and for knife use. Among both men and women, left-handed writers were significantly more likely to own a car, own a washing machine and have a bank account. Left-handed women (using both measures) had a higher income than right-handed women. Among men, left-handers for knife use had a higher income than right-handers. The results of our study suggest that the previously observed socioeconomic advantage of left-handers in Western populations also applies to non-Western populations, at least in the urban environment studied. However, we did not detect any difference in the number of children. We discussed how the frequency-dependent socioeconomic status advantage could be responsible for the persistence of left-handers throughout human evolution.     

Alteration of period and amplitude of circadian rhythms in shift workers. With special reference to temperature, right and left hand grip strength

48 male shift workers in various industries volunteered to document circadian rhythms in sleeping and working, oral temperature, grip strength of both hands, peak expiratory flow and heart rate. All physiological variables were self-measured 4 to 5 times a day for 2 to 4 weeks. Individual time series were analyzed according to several statistical methods (power spectrum, cosinor, chi squares, ANOVA, correlation, etc.) in order to estimate rhythm parameters such as circadian period (tau) and amplitude (A), and to evaluate subgroup differences with regard to tolerance to shift work, age, duration of shift work, speed of rotation and type of industry. The present study confirms for oral temperature and extends to other variables (grip strength of both hands, heart rate) that intolerance to shift work is frequently associated with both internal desynchronization and small circadian amplitude. The internal desynchronization among several circadian rhythms supports the hypothesis that these latter are driven by several oscillators. Many differences were observed between circadian rhythms in right and left hand grip strength: circadian tau in oral temperature was correlated with that in the grip strength of the dominant hand but not with that of the other hand; changes in tau s of the non-dominant hand were age-related but did not correlate with temperature tau; only the circadian A of the non-dominant hand was associated with a desynchronization. Thus, circadian rhythms in oral temperature and dominant hand grip strength may be driven by the same oscillator while that of the non-dominant hand may be governed by a different one. Internal desynchronization between both hand grip rhythms as well as desynchronization of performance rhythms reported by others provide indirect evidence that circadian oscillator(s) may be located in the human cerebral cortex.     

Dextrals and sinistrals (right-handers and left-handers): specificity of interhemispheric brain asymmetry and EEG coherence parameters

Data of literature about morphological, functional and biochemical specificity of the brain interhemispheric asymmetry of healthy right-handers and left-handers and about peculiarity of dynamics of cerebral pathology in patients with different individual asymmetry profiles are presented at the present article. Results of our investigation by using coherence parameters of electroencephalogram (EEG) in healthy right-handers and left-handers in state of rest, during functional tests and sleeping and in patients with different forms of the brain organic damage were analyzed too. EEG coherence analysis revealed the reciprocal changing of alpha-beta and theta-delta spectral bands in right-handers whilein left-handers synchronous changing of all EEG spectral bands were observed. Data about regional-frequent specificity of EEG coherence, peculiarity of EEG asymmetry in right-handers and left-handers, aslo about specificity of EEG spectral band genesis and point of view about a role of the brain regulator systems in forming of interhemispheric asymmetry in different functional states allowed to propose the conception about principle of interhermispheric brain asymmetry formation in left-handers and left-handers. Following this conception in dextrals elements of concurrent (summary-reciprocal) cooperation are predominant at the character of interhemispheric and cortical-subcortical interaction while in sinistrals a principle of concordance (supplementary) is preferable. These peculiarities the brain organization determine, from the first side, the quicker revovery of functions damaged after cranio-cerebral trauma in left-handers in comparison right-handers and from the other side - they determine the forming of the more expressed pathology in the remote terms after exposure the low dose of radiation.     

Visual Experience Determines the Use of External Reference Frames in Joint Action Control

Vision plays a crucial role in human interaction by facilitating the coordination of one''s own actions with those of others in space and time. While previous findings have demonstrated that vision determines the default use of reference frames, little is known about the role of visual experience in coding action-space during joint action. Here, we tested if and how visual experience influences the use of reference frames in joint action control. Dyads of congenitally-blind, blindfolded-sighted, and seeing individuals took part in an auditory version of the social Simon task, which required each participant to respond to one of two sounds presented to the left or right of both participants. To disentangle the contribution of external—agent-based and response-based—reference frames during joint action, participants performed the task with their respective response (right) hands uncrossed or crossed over one another. Although the location of the auditory stimulus was completely task-irrelevant, participants responded overall faster when the stimulus location spatially corresponded to the required response side than when they were spatially non-corresponding: a phenomenon known as the social Simon effect (SSE). In sighted participants, the SSE occurred irrespective of whether hands were crossed or uncrossed, suggesting the use of external, response-based reference frames. Congenitally-blind participants also showed an SSE, but only with uncrossed hands. We argue that congenitally-blind people use both agent-based and response-based reference frames resulting in conflicting spatial information when hands are crossed and, thus, canceling out the SSE. These results imply that joint action control functions on the basis of external reference frames independent of the presence or (transient/permanent) absence of vision. However, the type of external reference frames used for organizing motor control in joint action seems to be determined by visual experience.     

From a Somatotopic to a Spatiotopic Frame of Reference for the Localization of Nociceptive Stimuli

To react efficiently to potentially threatening stimuli, we have to be able to localize these stimuli in space. In daily life we are constantly moving so that our limbs can be positioned at the opposite side of space. Therefore, a somatotopic frame of reference is insufficient to localize nociceptive stimuli. Here we investigated whether nociceptive stimuli are mapped into a spatiotopic frame of reference, and more specifically a peripersonal frame of reference, which takes into account the position of the body limbs in external space, as well as the occurrence of external objects presented near the body. Two temporal order judgment (TOJ) experiments were conducted, during which participants had to decide which of two nociceptive stimuli, one applied to either hand, had been presented first while their hands were either uncrossed or crossed over the body midline. The occurrence of the nociceptive stimuli was cued by uninformative visual cues. We found that the visual cues prioritized the perception of nociceptive stimuli applied to the hand laying in the cued side of space, irrespective of posture. Moreover, the influence of the cues was smaller when they were presented far in front of participants’ hands as compared to when they were presented in close proximity. Finally, participants’ temporal sensitivity was reduced by changing posture. These findings are compatible with the existence of a peripersonal frame of reference for the localization of nociceptive stimuli. This allows for the construction of a stable representation of our body and the space closely surrounding our body, enabling a quick and efficient reaction to potential physical threats.     

"Biological geometry perception": visual discrimination of eccentricity is related to individual motor preferences

Background

In the continuum between a stroke and a circle including all possible ellipses, some eccentricities seem more “biologically preferred” than others by the motor system, probably because they imply less demanding coordination patterns. Based on the idea that biological motion perception relies on knowledge of the laws that govern the motor system, we investigated whether motorically preferential and non-preferential eccentricities are visually discriminated differently. In contrast with previous studies that were interested in the effect of kinematic/time features of movements on their visual perception, we focused on geometric/spatial features, and therefore used a static visual display.

Methodology/Principal Findings

In a dual-task paradigm, participants visually discriminated 13 static ellipses of various eccentricities while performing a finger-thumb opposition sequence with either the dominant or the non-dominant hand. Our assumption was that because the movements used to trace ellipses are strongly lateralized, a motor task performed with the dominant hand should affect the simultaneous visual discrimination more strongly. We found that visual discrimination was not affected when the motor task was performed by the non-dominant hand. Conversely, it was impaired when the motor task was performed with the dominant hand, but only for the ellipses that we defined as preferred by the motor system, based on an assessment of individual preferences during an independent graphomotor task.

Conclusions/Significance

Visual discrimination of ellipses depends on the state of the motor neural networks controlling the dominant hand, but only when their eccentricity is “biologically preferred”. Importantly, this effect emerges on the basis of a static display, suggesting that what we call “biological geometry”, i.e., geometric features resulting from preferential movements is relevant information for the visual processing of bidimensional shapes.