Left-Handers Are Less Lateralized Than Right-Handers for Both Left and Right Hemispheric Functions.

Many neuroscientific techniques have revealed that more left- than right-handers will have unusual cerebral asymmetries for language. After the original emphasis on frequency in the aphasia and epilepsy literatures, most neuropsychology, and neuroimaging efforts rely on estimates of central tendency to compare these two handedness groups on any given measure of asymmetry. The inevitable reduction in mean lateralization in the left-handed group is often postulated as being due to reversed asymmetry in a small subset of them, but it could also be due to a reduced asymmetry in many of the left-handers. These two possibilities have hugely different theoretical interpretations. Using functional magnetic resonance imaging localizer paradigms, we matched left- and right-handers for hemispheric dominance across four functions (verbal fluency, face perception, body perception, and scene perception). We then compared the degree of dominance between the two handedness groups for each of these four measures, conducting t-tests on the mean laterality indices. The results demonstrate that left-handers with typical cerebral asymmetries are less lateralized for language, faces, and bodies than their right-handed counterparts. These results are difficult to reconcile with current theories of language asymmetry or of handedness.

The depth and breadth of multiple perceptual asymmetries in right handers and non-right handers.

Several non-verbal perceptual and attentional processes have been linked with specialization of the right cerebral hemisphere. Given that most people have a left hemispheric specialization for language, it is tempting to assume that functions of these two classes of dominance are related. Unfortunately, such models of complementarity are notoriously hard to test. Here we suggest a method which compares frequency of a particular perceptual asymmetry with known frequencies of left hemispheric language dominance in right-handed and non-right handed groups. We illustrate this idea using the greyscales and colourscales tasks, chimeric faces, emotional dichotic listening, and a consonant-vowel dichotic listening task. Results show a substantial "breadth" of leftward bias on the right hemispheric tasks and rightward bias on verbal dichotic listening. Right handers and non-right handers did not differ in terms of proportions of people who were left biased for greyscales/colourscales. Support for reduced typical biases in non-right handers was found for chimeric faces and for CV dichotic listening. Results are discussed in terms of complementary theories of cerebral asymmetries, and how this type of method could be used to create a taxonomy of lateralized functions, each categorized as related to speech and language dominance, or not.

Do left hand reaction time advantages depend on localising unpredictable targets?

Asymmetries in hand movements have routinely been attributed to properties of the two cerebral hemispheres. In right-handed participants, the non-dominant left hand tends to have shorter reaction times, with the dominant right hand achieving shorter movement durations as well as higher peak velocities. The root cause of the surprising left hand RT effect has been debated, largely in the context of right hemisphere specialisation in attention, visuospatial abilities, or "premotor" processes. Mieschke et al. (Brain Cognit 45:1, 2001) and Barthélémy and Boulinguez ( Behav Brain Res 133:1, 2002) both tried to dissociate "premotor" processes explaining the left hand RT advantage, using reaching paradigms where at least one condition required target detection, but no visually guided aiming movement. Unfortunately, the studies obtained conflicting results and conclusions. In the present study, we attempted to re-examine this kind of paradigm with methodological improvements, such as using a task with higher visuospatial demands. Our results demonstrate that whilst RTs are longer as movement complexity increases across three conditions, the left hand RT advantage is present across all conditions-and no significant interaction between hand and condition was found. No significant hand differences were found in peak velocity or duration. These results suggest that the left hand RT advantage cannot be due to movement planning advantages of the right hemisphere, and instead should be attributed to sustained attention/vigilance lateralisation to the right cerebral hemisphere.

Hemispheric asymmetry of hand and tool perception in left- and right-handers with known language dominance.

Regions in the brain that are selective for images of hands and tools have been suggested to be lateralised to the left hemisphere of right-handed individuals. In left-handers, many functions related to tool use or tool pantomime may also depend more on the left hemisphere. This result seems surprising, given that the dominant hand of these individuals is controlled by the right hemisphere. One explanation is that the left hemisphere is dominant for speech and language in the majority of left-handers, suggesting a supraordinate control system for complex motor sequencing that is required for skilled tool use, as well as for speech. In the present study, we examine if this left-hemispheric specialisation extends to perception of hands and tools in left- and right-handed individuals. We, crucially, also include a group of left-handers with right-hemispheric language dominance to examine their asymmetry biases. The results suggest that tools lateralise to the left hemisphere in most right-handed individuals with left-hemispheric language dominance. Tools also lateralise to the language dominant hemisphere in right-hemispheric language dominant left-handers, but the result for left-hemispheric language dominant left-handers are more varied, and no clear bias towards one hemisphere is found. Hands did not show a group-level asymmetry pattern in any of the groups. These results suggest a more complex picture regarding hemispheric overlap of hand and tool representations, and that visual appearance of tools may be driven in part by both language dominance and the hemisphere which controls the motor-dominant hand.

Hemifield or hemispace: what accounts for the ipsilateral advantages in visually guided aiming?

Aiming movements to targets presented on the same side as the reaching limb are faster and more accurate than movements made across the body. These advantages are typically attributed to within-hemisphere sensorimotor control. However, contrary to the within- versus between-hemisphere model, we have shown that some of these advantages tend to go with the side of the movement, rather than the side of the target (Carey et al. Exp Brain Res 112:496-504, 1996; Carey and Otto-de Haart Neuropsychologia 39:894, 2001). Barthélémy and Boulinghez (Exp Brain Res 147:305-312, 2002) acknowledge that our biomechanical account fits data for post-onset movement parameters such as peak velocity and duration, yet they report evidence for some within- versus between-hemisphere contributions to reaction time (RT) advantages. To examine a possible difference between early and late movement kinematics fitting these alternative models, we have dissociated field and space in a different way, which required arm movements with differential inertial consequences, as well as unpredictability of target location in terms of visual field. The data suggest that visual field may contribute some of the variance to hemispatial effects, but only for the right hand. In a second experiment, we used an antipointing task to examine hemispatial versus visual field effects on RTs and to revisit the possible hand difference identified in experiment 1. We found that hemispace accounted for all of the ipsilateral advantages, including RT, for both right and left hands. Results are discussed in terms of the computational requirements of eye-hand coordination in relative unconstrained conditions.

Analysis of distributions reveals real differences on dichotic listening scores between left- and right-handers.

About 95% of right-handers and 70% of left-handers have a left-hemispheric specialization for language. Dichotic listening is often used as an indirect measure of this language asymmetry. However, while it reliably produces a right-ear advantage (REA), corresponding to the left-hemispheric specialization of language, it paradoxically often fails to obtain statistical evidence of mean differences between left- and right-handers. We hypothesized that non-normality of the underlying distributions might be in part responsible for the similarities in means. Here, we compare the mean ear advantage scores, and also contrast the distributions at multiple quantiles, in two large independent samples (Ns = 1,358 and 1,042) of right-handers and left-handers. Right-handers had an increased mean REA, and a larger proportion had an REA than in the left-handers. We also found that more left-handers are represented in the left-eared end of the distribution. These data suggest that subtle shifts in the distributions of DL scores for right- and left-handers may be at least partially responsible for the unreliability of significantly reduced mean REA in left-handers.

Inconsistent language lateralisation - Testing the dissociable language laterality hypothesis using behaviour and lateralised cerebral blood flow.

BACKGROUND: Most people have strong left-brain lateralisation for language, with a minority showing right- or bilateral language representation. On some receptive language tasks, however, lateralisation appears to be reduced or absent. This contrasting pattern raises the question of whether and how language laterality may fractionate within individuals. Building on our prior work, we postulated (a) that there can be dissociations in lateralisation of different components of language, and (b) these would be more common in left-handers. A subsidiary hypothesis was that laterality indices will cluster according to two underlying factors corresponding to whether they involve generation of words or sentences, versus receptive language. METHODS: We tested these predictions in two stages: At Step 1 an online laterality battery (Dichotic listening, Rhyme Decision and Word Comprehension) was given to 621 individuals (56% left-handers); At Step 2, functional transcranial Doppler ultrasound (fTCD) was used with 230 of these individuals (51% left-handers). 108 left-handers and 101 right-handers gave useable data on a battery of three language generation and three receptive language tasks. RESULTS: Neither the online nor fTCD measures supported the notion of a single language laterality factor. In general, for both online and fTCD measures, tests of language generation were left-lateralised. In contrast, the receptive tasks were at best weakly left-lateralised or, in the case of Word Comprehension, slightly right-lateralised. The online measures were only weakly correlated, if at all, with fTCD measures. Most of the fTCD measures had split-half reliabilities of at least .7, and showed a distinctive pattern of intercorrelation, supporting a modified two-factor model in which Phonological Decision (generation) and Sentence Decision (reception) loaded on both factors. The same factor structure fitted data from left- and right-handers, but mean scores on the two factors were lower (less left-lateralised) in left-handers. CONCLUSIONS: There are at least two factors influencing language lateralization in individuals, but they do not correspond neatly to language generation and comprehension. Future fMRI studies could help clarify how far they reflect activity in specific brain regions.

Gating of vibrotactile detection during visually guided bimanual reaches.

It is far more difficult to detect a small tactile stimulation on a finger that is moving compared to when it is static. This suppression of tactile information during motion, known as tactile gating, has been examined in some detail during single-joint movements. However, the existence and time course of this gating has yet to be examined during visually guided multi-joint reaches, where sensory feedback may be paramount. The current study demonstrated that neurologically intact humans are unable to detect a small vibratory stimulus on one of their index fingers during a bimanual reach toward visual targets. By parametrically altering the delay between the visual target onset and the vibration, it was demonstrated that this gating was even apparent before participants started moving. A follow up experiment using electromyography indicated that gating was likely to occur even before muscle activity had taken place. This unique demonstration of tactile gating during a task reliant on visual feedback supports the notion this phenomenon is due to a central command, rather than a masking of sensory signals by afferent processing during movement.

Bimanual reaching across the hemispace: which hand is yoked to which?

When both hands perform concurrent goal-directed reaches, they become yoked to one another. To investigate the direction of this coupling (i.e., which hand is yoked to which), the temporal dynamics of bimanual reaches were compared with equivalent-amplitude unimanual reaches. These reaches were to target pairs located on either the left or right sides of space; meaning that in the bimanual condition, one hand's contralateral (more difficult) reach accompanied by the other hand's ipsilateral (easier) reach. By comparing which hand's difficult reach was improved more by the presence of the other hand's easier ipsilateral reach, we were able to demonstrate asymmetries in the coupling. When the cost of bimanual reaching was controlled for the contralateral reaching left hand's performance was improved, suggesting that the left hand is yoked to the right during motor output. In contrast, the right hand showed the greatest improvements for contralateral reaching in terms of reaction time, pointing toward a dominant role for the left hand in the processes prior to movement onset. The results may point toward a mechanism for integrating the unitary system of attention with bimanual coordination.

The validity and reliability of quantifying hemispheric specialisation using fMRI: Evidence from left and right handers on three different cerebral asymmetries.

Neuroimaging has tremendous potential for quantifying hemispheric specializations. However, the possibilities remain under-utilized, in part, given some of the complexities in quantifying any differences in a reliable, transparent fashion. A second issue with hemispheric asymmetries is that they are extremely one-sided in most people. This skew limits the generalisability of any findings to those participants with rarer forms of cerebral asymmetry. Here, we demonstrate usefulness of an approach developed by Wilke and Lidzba, (J Neurosci Meth, 163, 2007), which allows for threshold-independent estimates of cerebral asymmetry to be calculated in individual participants. We compared these estimates from two separate runs for three different cerebral asymmetries in the same participants. We circumvented the skewed nature of this type of data in two ways; first, we scanned a large number of non-right handed participants, and second, we included asymmetries that favour the right hemisphere in right handers, which we had reason to believe were less skewed than those related to speech and language. Verbal fluency and two visuoperceptual asymmetries were localized in a sample of 33 right handed and 60 non-right handed participants. Laterality indices (LIs), which quantify the direction and strength of an asymmetry, were calculated for BOLD activity relating to language, face perception, and body perception in each run separately. Run 1 - run 2 correlations were all statistically significant and surprisingly sizeable (r = 0.89 to r = 0.62), considering the relatively short amount of time on task within our particular localizers. This noteworthy success validates a number of useful ways that functional neuroimaging can be used to advance understanding of cerebral asymmetries.

Four meta-analyses across 164 studies on atypical footedness prevalence and its relation to handedness.

Human lateral preferences, such as handedness and footedness, have interested researchers for decades due to their pronounced asymmetries at the population level. While there are good estimates on the prevalence of handedness in the population, there is no large-scale estimation on the prevalence of footedness. Furthermore, the relationship between footedness and handedness still remains elusive. Here, we conducted meta-analyses with four different classification systems for footedness on 145,135 individuals across 164 studies including new data from the ALSPAC cohort. The study aimed to determine a reliable point estimate of footedness, to study the association between footedness and handedness, and to investigate moderating factors influencing footedness. We showed that the prevalence of atypical footedness ranges between 12.10% using the most conservative criterion of left-footedness to 23.7% including all left- and mixed-footers as a single non-right category. As many as 60.1% of left-handers were left-footed whereas only 3.2% of right-handers were left-footed. Males were 4.1% more often non-right-footed compared to females. Individuals with psychiatric and neurodevelopmental disorders exhibited a higher prevalence of non-right-footedness. Furthermore, the presence of mixed-footedness was higher in children compared to adults and left-footedness was increased in athletes compared to the general population. Finally, we showed that footedness is only marginally influenced by cultural and social factors, which play a crucial role in the determination of handedness. Overall, this study provides new and useful reference data for laterality research. Furthermore, the data suggest that footedness is a valuable phenotype for the study of lateral motor biases, its underlying genetics and neurodevelopment.

The bi-pedal ape: plasticity and asymmetry in footedness.

A strong preference for using the right foot for skilled activities parallels a similar side bias for hand use. However, many neuropsychologists, sports scientists and sports commentators argue that right-foot bias in soccer is reduced or even eliminated by practice. This sort of plasticity is an important component of the principle genetic theories of handedness, yet very little is known about the relative asymmetries in the many unipedal skills required in soccer at amateur or professional levels. The first study examined self-report of hand and foot bias in a sample (n=400) of amateur soccer players, in addition to information about their practice with the non-preferred foot. A second study quantified foot use on the pitch in a large sample (n=426) of professional soccer players. The majority of the amateurs reported a right-foot bias that is very similar to that seen in the general population ( approximately 80%). However they only endorse strong biases for "closed" (self-paced) soccer actions like penalty and other free kicks, which give players unlimited time and space for preparation and execution. Although there was a very slight tendency for less right-foot bias in the professionals ( approximately 75%), as assessed by actual foot use rather than questionnaire, few players show anything like what could be described as two-footed play. This bias, unlike in the reports of amateurs, were for all of the behaviours investigated, not just so-called "skilled" behaviours. Finally, when outcomes of preferred and non-preferred foot behaviours were contrasted, the professionals were remarkably adept on those rare occasions when they use their non-preferred foot, suggesting that skill cannot explain asymmetry of choice. These results are discussed in terms of 1) limitations of self-report on questionnaires for predicting actual on the field behaviour, 2) the surprising absence of plasticity in foot use, given the importance of learning, experience and culture in models of handedness and footedness, and 3) a left hemisphere lateralised intentional system as important for the selection of movements as for their execution.

Rightward biases during bimanual reaching.

Two experiments were carried out to investigate whether attention is biased toward the right hand of right handers during bimanual coordination (Peters 1981). A novel discontinuous double-step reaching task was developed, where right-handed participants executed a bimanual reach followed by a left or right hand unimanual reach. Asymmetries in the downtime between the bimanual and unimanual reach portions (the refractory period) were used to infer the direction of attention. A shorter right hand refractory period was found in the first experiment, indicating a rightward bias in attention. In a second experiment, shifting the focus of attention during the bimanual portion of the reach altered the direction and magnitude of the asymmetry in a way consistent with the attentional bias hypothesis. The role of attention during bimanual reaching, and a further programme of experimental work aimed at clarifying the nature of these rightward biases during discrete bimanual coordination is discussed.

Does localisation blindsight extend to two-dimensional targets?

Residual sensorimotor skills which survive compromise of the geniculostriate visual system may depend on activity of the dorsal stream of extrastriate occipitoparietal cortex. These circuits are crucial for controlling hand and eye movements to targets in a three-dimensional world. Remarkably, demonstrations of above chance localisation by hand and by eye in blindsight patients have used luminous targets that were only varied in one spatial dimension. These limitations result in experimental confounds. In the present study we examined saccadic and manual localisation in a well-studied patient (DB) to positions that were varied in 1 or 2 dimensions, using targets which control for luminance artefacts. We found that his good manual localisation without awareness in 1D conditions was relatively preserved when the targets were varied in 2D. In stark contrast, saccadic performance was completely attenuated with 2D targets. These paradoxical results are difficult to reconcile with feedforward models of eye-hand coordination and with accounts of localisation that depend on intact multidimensional representations of the visual fields in non-geniculostriate systems.

Investigating bimanual coordination in dominant and non-dominant virtual hands.

A bias in attention towards the dominant hand has been cited as a possible factor in the lateralisation of human bimanual coordination (Peters, 1981). A mirror was placed between the hands of 18 dextral participants performing rhythmic anti-phase movements. This set-up gave the appearance of a reflected virtual hand (moving in time with the un-occluded hand), in the same spatial location as the occluded left or right hand. This asymmetrical conflict between vision and action examined whether the left hand would show higher levels of error when replaced by a virtual right hand than the converse condition. Higher levels of error were observed during performance of the anti-phase pattern overall in the conditions where the mirror was present (compared to control conditions without the mirror). However, this effect did not differ between hands. The implications for the mirror paradigm, possible explanations for the lack of asymmetry, and the consequences for the attentional bias hypothesis are discussed.

Visuomotor 'immunity' to perceptual illusion: a mismatch of attentional demands cannot explain the perception-action dissociation.

Recent findings of visuomotor immunity to perceptual illusions have been attributed to a perception-action division of labour within two anatomically segregated streams in the visual cortex. However, critics argue that such experimental findings are not valid and have suggested that the perception-action dissociations can be explained away by differential attentional/processing demands, rather than a functional dissociation in the neurologically intact brain: perceptual tasks require processing of the entire illusion display while visuomotor tasks only require processing the target that is acted upon. The present study examined whether grasping of the Müller-Lyer display would remain immune to the illusion when the task required the direction of attention or a related resource towards both Müller-Lyer shafts. Twelve participants were required to match and grasp two Müller-Lyer shafts bimanually (i.e. one with each hand). It was found that bimanual grasping was not significantly affected by the illusion, while there was a highly significant illusion effect on perceptual estimation by matching. Furthermore, it was established that this dissociation did not result from a differing baseline rate of change in manual estimation and grasping aperture to a change in physical object size. These findings provide further support for the postulated perception-action dissociation and fail to uphold the idea that grasping 'immunity' to the Müller-Lyer illusions merely represents an experimental artefact.

Tapping, grasping and aiming in ideomotor apraxia.

Very few studies have investigated sensorimotor control in apraxia using tasks that differ in movement complexity. Nevertheless, there is some evidence to suggest that spontaneous behaviour, although relatively preserved, can be rather clumsy or awkward, and that patients with ideomotor apraxia may have subtle kinematic abnormalities in movements made in the laboratory. It remains unclear whether patients with ideomotor apraxia perform normally on movements such as visually guided aiming, that may not depend on higher-order, more cognitive, processes and that are relatively unguided by overlearned contexts. In this study, three different sensorimotor tasks were given to the same sample of patients with quantified apraxic disturbance. Finger tapping, goal-directed grasping and aiming with and without visual feedback were examined in these patients. A clear dissociation was found between grossly impaired gesture imitation and intact motor programming of goal-directed movements with visual feedback. Apraxic patients were, however, impaired on aiming movements without visual feedback, suggesting that apraxia is associated with an increased reliance on integration of online visual information with feedforward/feedback somatosensory and motor signals. Furthermore, patients were impaired on single finger tapping which was a surprisingly good predictor of apraxia severity.

Pointing to places and spaces in a patient with visual form agnosia.

Previous investigations of visuospatial abilities in the visual form agnosic patient D.F. suggest that her egocentric sensorimotor processing is intact while her 'allocentric' judgments of spatial position are impaired. The current investigation extends these previous observations by comparing D.F.'s performance at pointing to a set of spatially distributed stimuli, either directly or by 'pantomiming' the responses in an adjacent homologous workspace. The results showed accurate sensorimotor localization when D.F. pointed directly to single targets or to sequences of targets, presumably as she could use egocentric visual coding. In spite of making relatively spared spatial judgments about the arrays, however, D.F. performed quite poorly when copying them and on the pantomimed pointing task. In this latter task good performance presumably depends on an ability to represent both the categorical and coordinate properties of the array (as does copying them), and to translate these into the effector-based coordinates required for accurate action. D.F.'s pantomimed pointing was similar to her copies of target arrays, as in both tasks there was evidence of spared (although somewhat degraded) appreciation of the relative spatial positions of the stimuli. Remarkably, her accuracy in this allocentric task was not worsened by longer pointing sequences. It is possible that D.F.'s degraded performance reflects a relative (though not complete) preservation of categorical coding within the ventral stream, despite a loss of coordinate coding there.

Are there right hemisphere contributions to visually-guided movement? Manipulating left hand reaction time advantages in dextrals.

Many studies have argued for distinct but complementary contributions from each hemisphere in the control of movements to visual targets. Investigators have attempted to extend observations from patients with unilateral left- and right-hemisphere damage, to those using neurologically-intact participants, by assuming that each hand has privileged access to the contralateral hemisphere. Previous attempts to illustrate right hemispheric contributions to the control of aiming have focussed on increasing the spatial demands of an aiming task, to attenuate the typical right hand advantages, to try to enhance a left hand reaction time advantage in right-handed participants. These early attempts have not been successful. The present study circumnavigates some of the theoretical and methodological difficulties of some of the earlier experiments, by using three different tasks linked directly to specialized functions of the right hemisphere: bisecting, the gap effect, and visuospatial localization. None of these tasks were effective in reducing the magnitude of left hand reaction time advantages in right handers. Results are discussed in terms of alternatives to right hemispheric functional explanations of the effect, the one-dimensional nature of our target arrays, power and precision given the size of the left hand RT effect, and the utility of examining the proportions of participants who show these effects, rather than exclusive reliance on measures of central tendency and their associated null hypothesis significance tests.

Processing biases towards the preferred hand: valid and invalid cueing of left- versus right-hand movements.

A Posner-like paradigm was employed to investigate the effects of valid and invalid cueing of each hand on reaction time, movement time and peak velocity in an aiming task. Given claims of left hemisphere superiority in movement selection and inhibition (and the privileged within-hemisphere access of the right hand to such systems), it was hypothesised that invalidly cueing the left hand (i.e. right-hand movement precued, left-handed movement required by a go signal) would result in increased reaction time relative to invalid right-hand cueing. The hypothesis was not confirmed as reaction times of both hands were slowed equivalently by invalid cueing. Nevertheless, it was found that the movement duration of the left hand was increased substantially by invalid cueing, while the right hand was unaffected on this measure, suggesting a possible intentional rather than attentional difference between the two hands. These results are discussed in terms of a possible asymmetry of intentional processes related to hand movement and the right-hand advantage in movement duration.