Interhemispheric Cooperation of Left- and Righthanders in Mental Calculation Tasks

Relations between handedness and interhemispheric processing in cognitive tasks were examined. Thirty six right-handers and thirty left-handers (familial and nonfamilial left-handers) were asked to add two numbers which were presented tachistoscopically. Two numbers were displayed either to one visual field, or one to the left and one to the right visual field simultaneously. In Experiment 1, the numbers were displayed in Arabic numerals, and in Experiment 2 one of the numbers was displayed in Kanji and one in Arabic numerals. The results of Experiment 1 showed a bilateral advantage, however no subject group difference was shown in the performance of the three (left unilateral, right unilateral, and bilateral) presentation conditions. Though familial left-handers showed a weaker tendency to different patterns than right and non-familial left-handers, the results of Experiment 2 were largely similar to those of Experiment 1. These findings suggest that interhemispheric processing may not differ between left- and righthanders.     

Temporal integration of form as a function of subject handedness and retinal locus of presentation

Two experiments are reported examining differences in perceptual processing as a function of subject handedness. Experiment 1 compared performance in the left vs right visual fields of right- vs left-handed subjects. Only right-handed subjects exhibited an interaction of visual field with spatial frequency (as mediated by interflash interval), suggesting that left-handers are not differentially lateralized for high vs low spatial frequency processing. Experiment 2 examined foveal performance in left-handers vs right-handers with or without familial sinistrality. Subjects with personal or familial sinistrality exhibited superior performance when the processing of low frequencies was required, suggesting that sinistrality may confer an advantage in the processing of lower spatial frequencies.     

Bilateral processing of redundant information: the influence of stimulus notation and processing speed in number comparison

Improved performance is generally observed when identical copies of a stimulus are presented bilaterally to both visual half fields (VHFs), relative to unilateral single presentations. We investigated the influence of stimulus notation and processing speed on this bilateral redundancy gain (B-RG) in number comparison. Two experiments were performed with consistent (i.e., two identical copies of Arabic digits or word numerals) and inconsistent (i.e., an Arabic digit in one VHF with a word numeral of the same value in the other VHF) displays (see also Marks and Hellige, 2003 for a similar design in number naming). In Experiment 1, the processing speeds of the Arabic digits were faster than the ones of the word numerals while in Experiment 2, the processing speeds of the two stimulus notations were kept equal. A significant difference between consistent and inconsistent conditions was restricted to Experiment 1 in which the processing speeds of the two stimulus notations diverged. These results are discussed in the framework of a simple race model in which two independent hemispheric processes compete for the control of response, and the faster process wins.     

Unconscious processing of Arabic numerals in unilateral neglect

This study explores the processing of Arabic numerals in three patients with dense left unilateral neglect. Three tasks have been used: a test of visual awareness (is the stimulus on the left, on the right or on both sides?), a number comparison task (is the number larger or smaller than 5?), a number parity judgment task (is the number odd or even?). The test of visual awareness indicated that all three patients were completely unaware of the stimuli presented in the left hemifield. Despite this, the number comparison and number parity judgment tasks clearly indicated that Arabic numerals were unconsciously processed at semantic level (i.e. quantity). These results show that patients with left unilateral neglect can still semantically process Arabic numerals unconsciously.     

Bilateral processing of numbers: parallel processing versus functional unity

Thirty-six normal subjects had to add two numbers ranging from 3 to 9 which were presented tachistoscopically. The two numbers were displayed either unilaterally to the left or right visual field, or bilaterally with one number to the left and one to the right visual field. They were represented as two word names, two dot clusters, or two bargraphs, or the two numbers were represented inconsistently in two different of these representations. The results showed that bilateral processing of a word and a bargraph was performed without visible interactions, and the hemispheres performed independently. When the display included a dot cluster, however, its processing (sequential vs. holistic) appeared to depend on the type of stimulus processed by the contralateral hemisphere, indicating strong interhemispheric interactions.     

Hemispheric dominance for speech and calculation: electrophysiologic correlates of left dominance in left handedness

Slow brain potentials were recorded in left-handers and right-handers during: (i) processing of language and mental arithmetic tasks, without vocalization, and (ii) subsequent writing down of the answers with either the right or left hand. Left-sided laterality of negative potentials was taken as evidence of hemispheric dominance. It appeared during the processing of words and numbers in 26 of the 30 left-handers and was localized mainly in the left frontal and temporal parietal regions. Similar results were found with the right-handers. This electrophysiological evidence indicates that the left hemisphere is dominant for language and calculation in the vast majority of left-handers. Only when writing with either their left or right hand do left-handers show less left-sided laterality than right-handers.     

Zur hemispha¨rendominanz fu¨r sprache und rechnen: Elektrophysiologische korrelate einer linksdominanz bei linksha¨ndern

Slow brain potentials were recorded in left-handers and right-handers during: (i) processing of language and mental arithmetic tasks, without vocalization, and (ii) subsequent writing down of the answers with either the right or left hand. Left-sided laterality of negative potentials was taken as evidence of hemispheric dominance. It appeared during the processing of words and numbers in 26 of the 30 left-handers and was localized mainly in the left frontal and temporal parietal regions. Similar results were found with the right-handers. This electrophysiological evidence indicates that the left hemisphere is dominant for language and calculation in the vast majority of left-handers. Only when writing with either their left or right hand do left-handers show less left-sided laterality than right-handers.     

An analysis of the naming deficit of left-handers

Left-handers have been found to have a naming deficit when confronted with briefly flashed words. Three alternative explanations of this deficit were tested: whether it was due to deviant interhemispheric cooperation, relatively diffuse neural organization, or left hemispheric dysfunction. In Experiment I, four words were presented for 190 ms, unilaterally (to a single hemisphere), or bilaterally (between the hemispheres). Although left-handers (N = 48) named significantly less than right-handers (N = 30), performance was parallel across hemispheric conditions. Experiment II required semantic categorization, but not naming. The performance of the left-handers (N = 27) was indistinguishable from that of right-handers (N = 27) both in terms of overall performance and interhemispheric collaboration. It is concluded that the linguistic deficit of left-handers is specific to oral naming and that it is not caused by deviant interhemispheric cooperation.     

Bilateral field interactions and hemispheric asymmetry in number comparison

We investigated bilateral field interactions and hemispheric asymmetry in number comparison. A numerical comparison task with three different stimulus notations (Arabic numerals, word numerals and bar graphs) was used. In all conditions, a target was displayed in one visual field, simultaneously with a distractor of the same format in the other visual field. Participants had to indicate manually whether the magnitude of the target was small or large, ignoring the distractor stimulus. Only in the condition with Arabic numerals did we obtain some evidence for a LVF advantage, which argues against a strong laterality of number magnitude representations. Significant interactions between target and distractor values were observed, indicating rich interhemispheric interactions. The interactions were mainly situated at the response stage, but the presence of a bilateral identity effect also points to interactions at the input level.     

Tachistoscopic recognition of Kana and Hangul words,handedness and shift of laterality difference

Nonsense Kana words as verbal stimuli and Hangul words (Korean orthographic characters) as non-verbal were presented tachistoscopically in the left or right visual field to normal right-handed, non-familial left-handed and familial left-handed Japanese subjects, who have not seen Hangul characters. Right-handers showed a significant right (or left) field superiority for the recognition of verbal (or non-verbal) stimuli. This pattern of differences found in right-handers is, though to a slightly lesser degree, also present in non-familial left-handers, while it is absent in familial left-handers. Initial left field superiority for Hangul word recognition shifted to no laterality difference in left-handers as well as in right-handers during the learning period for Hangul words.     

Motor control of the hands: the effect of familial sinistrality.

Left- and right-handed subjects, selected on the basis of degree of hand preference and for the presence or absence of familial sinistrality, responded to monaurally presented tonal stimuli (440 Hz note played on four different instruments) using their right and left hands on separate occasions. It was found that in both the strong left-handers and the inconsistent strong right-handers, motor control of the hands was related to familial sinistrality (FS). Specifically, strong left-handers and inconsistent strong right-handers with FS have a difference in the motor control of the hands in the left hemisphere, with a left hemisphere-left hand advantage. Strong left-handers and inconsistent strong right-handers with no FS have a difference in the motor control of the hands in the right hemisphere, with a right hemisphere-left hand advantage.     

Avowed, assessed, and familial handedness and differential hemispheric processing of brief sequential and non-sequential visual stimuli

Right-handers with no family history of left-handedness (RHFH-), right-handers with such history (RHFH+), and left-handers (LH) were tested on a lateralized letter masking task. Only the RHFH - group showed significant right visual field superiority on the masking task. The results would seem to reflect interhemispheric communication efficiency rather than which hemisphere is dominant for language expression.     

Interhemispheric cooperation and activation in integration of verbal information

Subjects had to judge the semantic relation between two tachistoscopically presented German adjectives. In Exp. I, the two words were projected simultaneously, either both to the left visual field (i.e. to the right hemisphere, LVF-RH), or both to the right visual field (i.e. to the left hemisphere, RVF-LH), or one word to each visual field (i.e. to both hemispheres, EVF-BH). The results yielded a significantly higher performance in the condition EVF-BH than in the two unilateral conditions, without a significant difference between the latter two conditions. In Exp. II, the two words were shown successively. With bilateral presentation, a distinction was made as to whether the left visual field (EVF-BH-L) or the right visual field (EVF-BH-R) was stimulated first. The significantly highest performance was shown in conditions RVF-LH and EVF-BH-R compared with the condition EVF-BH-L, and the significantly poorest performance in the condition LVF-RH compared with the condition EVF-BH-L. The results provide evidence for (1) an intrahemispheric interference with simultaneous unilateral presentation, (2) a cooperative interhemispheric interaction with simultaneous bilateral presentation, and (3) an interhemispheric priming effect with successive bilateral presentation when the left hemisphere was stimulated first.     

Bilateral visual field processing and evoked potential interhemispheric transmission time

The relationship between the efficiency of interhemispheric interactions via the corpus callosum and the speed and accuracy in making comparisons of information simultaneously presented to the right and left visual fields was studied by comparing bilateral (vs unilateral) advantages in matching letters, with evoked potential measures of interhemispheric transmission time (EP-IHTT). The primary findings was a strong correlation suggesting that larger bilateral field advantages in reaction time are associated with faster EP-IHTT. However, the association between EP-IHTT and bilateral advantage was strong only for transmission speed from left hemisphere to right hemisphere, but not for speed of transmission in the opposite direction. The data are consistent with a hypothesis of asymmetric homologue enhancement, i.e. a directionally asymmetric callosal influence which facilities processing of letter stimuli in the right hemisphere allowing for increased response speed and accuracy of bilateral visual field comparisons.     

Interhemispheric connectivity during lateralized lexical decision

The well‐established right visual field (RVF‐lh) advantage in word recognition is commonly attributed to the typical left hemisphere dominance in language; words presented to the LVF‐rh are processed less efficiently due to the need for transcallosal transfer from the right to left hemisphere. The exact stage for this hemispheric transfer is currently unsettled. Some studies suggest that transfer occurs at very early stages between primary visual regions, whereas other studies suggest that transfer occurs between the left visual word form area and its right hemisphere homolog. This study explores these conflicting accounts and finds evidence for both. Participants conducted a lateralized lexical decision task with both unilateral and bilateral display conditions. Connectivity analyses were conducted from magnetoencephalography signals that were localized to the left middle occipital gyrus (LMOG), right middle occipital gyrus (RMOG), left visual word form area (LVWFA), and right visual word form area (RVWA). Results from unilateral trials showed asymmetrical interhemispheric connectivity from the RMOG to LMOG and symmetrical interhemispheric connectivity between the LVWFA and RVWFA. Furthermore, bilateral presentations led to reduced interhemispheric connectivity between both homologous region of interest pairs. Together, these results suggest that lateralized word recognition involves multiple stages of interhemispheric interactions and that these interactions are reduced with bilateral displays.     

Hemispheric differences between left and right number representations: effects of conscious and unconscious priming

The contribution of each hemisphere to the generation of number representations was investigated by two lateralized priming experiments in which participants had to compare Arabic digits to a fixed standard of four. In Experiment 1, unmasked primes (Arabic digits or word numerals) were used. In Experiment 2, masked primes were presented consciously or subconsciously. In both experiments similar priming effects were found in the left (LH) and the right hemisphere (RH) when the prime was presented consciously. However, asymmetries emerged when the primes were presented subconsciously: while the priming effects of digits and word numbers were equally large in the right visual half field (RVF-LH), the influence of the word prime on the semantic and the response stage of the left visual half field (LVF-RH) was absent, indicating that a word prime was no longer processed when it was presented subconsciously in LVF-RH. We believe that the origin of the latter effect can be attributed to a failure to transfer word number primes from the RH to the LH when attentional resources are restricted.     

Right hemispheric dominance and interhemispheric cooperation in gaze-triggered reflexive shift of attention

Aims: The neural substrate for the processing of gaze remains unknown. The aim of the present study was to clarify which hemisphere dominantly processes and whether bilateral hemispheres cooperate with each other in gaze‐triggered reflexive shift of attention. Methods: Twenty‐eight normal subjects were tested. The non‐predictive gaze cues were presented either in unilateral or bilateral visual fields. The subjects localized the target as soon as possible. Results: Reaction times (RT) were shorter when gaze‐cues were congruent toward than away from targets, whichever visual field they were presented in. RT were shorter in left than right visual field presentations. RT in mono‐directional bilateral presentations were shorter than both of those in left and right presentations. When bi‐directional bilateral cues were presented, RT were faster when valid cues were presented in the left than right visual fields. Conclusion: The right hemisphere appears to be dominant, and there is interhemispheric cooperation in gaze‐triggered reflexive shift of attention.     

Dyslexic adolescents: evidence of impaired visual and auditory language processing associated with normal lateralization and visual responsivity.

The questions of whether chronically dyslexic adolescent suffer any deficits of simple language stimulus processing or are less left hemisphere dominant than normal reading controls were addressed. The dyslexics were chosen for clarity of their specific reading problem and were older than dyslexics previously studied with lateralizing tests. Tasks administered in Experiment I were unilateral and bilateral tachistoscopic work recognitions and a tachistoscopic recognition report-time task for single lateralized letter stimuli. Experiment II, conducted a year later, readministered these tasks with modifications, and added dichotic digits and motor reaction time-stimulus detection tasks. It was concluded that right handed, chronic dyslexics: (1) possess left hemisphere language specialization; (2) show normal interhemispheric processing delays for single letter stimuli; (3) are, unlike nondyslexis but equally poor-reading Ss, clearly impaired in their efficiency of visual and auditory processing of simple language stimuli; (4) possess clear auditory memory deficits for verbal material; and (5) may possess an additional deficit of left hemisphere visual association area function.     

Influence of handedness on peripheral auditory asymmetry

It is well established that in humans many differences between right- and left-handers, anatomical, physiological and functional, exist. Left- and mixed-handedness is associated with greater bihemispheric representation of cognitive functions than in right-handers. Several studies indicate a left–right asymmetry in the function of hearing pathways between cochlea and auditory cortex, and furthermore, that this asymmetry is associated with handedness. Our investigation focuses on the medial olivo-cochlear system, which has been demonstrated to be more effective in the right than left ear in right-handers. The aim of the study was to investigate this auditory efferent system asymmetry according to handedness, gender, eyedness, footedness and the presence of spontaneous otoacoustic emissions. The medial efferent system has been found to be more effective in the right than left ear in right-handers, while functioning symmetrically in left-handers. Furthermore, the olivo-cochlear system, assumed to be involved in basic language processing, shows an asymmetrical pattern of functioning influenced by handedness as well as by hemispheric language representation. Reverse medial efferent system asymmetry was observed in left-handers compared to that in right-handers, on condition that only left-handed males were considered, or that the left-handers were also left-eyed, or that spontaneous otoacoustic emissions were present in the left ear of the left-handers, or when only left-handers without mixed-handers were considered.     

Handedness and the excitability of cortical inhibitory circuits

Inhibitory processes play a significant role in the control of goal-directed actions. To increase insights into these mechanisms as a function of handedness, we measured the transient inhibition of volitional motor activity induced by single pulse transcranial magnetic stimulation during bimanual isometric contractions with symmetrical and asymmetrical force demands. Here, we assess the cortical silent period (cSP), which associates with intrahemispheric inhibition, and the ipsilateral silent period (iSP), which provides an estimation of interhemispheric inhibition. The data showed that inhibitory processes support the functional regulation of bimanual motor output. Furthermore, right-handers demonstrated asymmetries in intra- and interhemispheric inhibition due to asymmetrical force requirements and hand dominance, whereas left-handers did not show marked differences. In particular, right-handers demonstrated increased inhibitory processing that favoured control of the dominant (left) hemisphere whereas both motor cortices exhibited equal capabilities in left-handers. These observations were specific to the bimanual nature of the task. The present results underline distinct organisational mechanisms of coordinated behaviour in right- and left-handers.