MRI Asymmetries of Broca''s Area: The Pars Triangularis and Pars Opercularis

Broca's area, which includes the pars triangularis (PTR) and pars opercularis (POP), is a neuroanatomic region important in speech-language production. Previous data demonstrated that PTR asymmetries are highly correlated with language dominance determined by selective hemispheric anesthesia or Wada testing, suggesting that asymmetries of the PTR may, in part, predict language dominance. The POP, however, has not been measured on volumetric magnetic resonance imaging (MRI), and therefore, it is unclear whether morphological asymmetries of the POP exist, and whether these asymmetries differ in right- and left-handers. The purpose of this study was to determine if measurable asymmetries of the POP exist on MRI, and whether the direction of the asymmetries differ in right- and left-handers. The PTR and POP were measured on volumetric MRI scans of 16 right-handers and 16 left-handers matched for age and gender. There was a significant leftward asymmetry of the PTR in right- and left-handers, although the asymmetry was reduced in the left-handers. In contrast, there was a leftward asymmetry of the POP in right-handers, and a rightward asymmetry in the left-handers. Handedness, derived from a handedness inventory, was positively correlated with POP asymmetry.     

Performance of subgroups of left-handers and right-handers

Fifty-three left-handers with consistent left-hand preferences (CLH), 65 left-handers with inconsistent hand preferences (ILH), and 57 right-handers (RH) were given unimanual and bimanual performance tests involving skill, speed, and strength as well as tests of articulatory speed and verbal fluency. Contrary to claims in the current literature (Ponton, 1987), CLHs and ILHs do not differ in quality and speed of performance, but, in some tests, they do show asymmetries in opposite directions. Thus, when left-handers are treated as a combined group, the faulty impression of a lack of between-hand asymmetries arises. The results suggest that a distinction between CLHs and ILHs yields subgroups with reliably different and distinctive performance patterns which are not trivially attributable to differences in strength of lateralization. CLHs behave much like mirror image RHs, whereas ILHs show a dissociation between strength, fine manual skill, attentional asymmetries.     

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

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

Subject variable effects in correlations between auditory and visual language processing asymmetries

The cross-modal correlations between auditory and visual language lateralization were examined as a function of the subject variables of handedness, sex, familial sinistrality, and handwriting posture. In this study, left-handers showed a significantly greater correlation between visual and auditory language processing asymmetries than right-handers, contradicting previous reports.     

Hand effects on mentally simulated reaching

Within the area of simulated (imagined) versus actual movement research, investigators have discovered that mentally simulated movements, like real actions, are controlled primarily by the hemispheres contralateral to the simulated limb. Furthermore, evidence points to a left-brain advantage for accuracy of simulated movements. With this information it could be suggested that, compared to left-handers, most right-handers would have an advantage. To test this hypothesis, strong right- and left-handers were compared on judgments of perceived reachability to visual targets lasting 150 ms in multiple locations of midline, right- and left-visual field (RVF/LVF). In reference to within group responses, we found no hemispheric or hand use advantage for right-handers. Although left-handers revealed no hemispheric advantage, there was a significant hand effect, favoring the non-dominant limb, most notably in LVF. This finding is explained in regard to a possible interference effect for left-handers, not shown for right-handers. Overall, left-handers displayed significantly more errors across hemispace. Therefore, it appears that when comparing hand groups, a left-hemisphere advantage favoring right-handers is plausible.     

Dichotic listening performance and writing posture in right- and left-handers

Right-handers and left-handers with the inverted (IN) and noninverted (NI) writing posture were tested on a dichotic consonant-vowel listening task and on two motor tasks (hand strength and speed of tapping). The results failed to show the differences between IN and NI right-handers reported by S. M. Tapley and M. P. Bryden (1983, Neuropsychologia, 21, 129-138) and there were no significant handedness x writing posture x ear interactions. A significant interaction between dichotic listening performance and writing posture was found; NI right-handers and IN left-handers had more correct responses and fewer intrusions than IN right-handers and NI left-handers. Left-handers and right-handers were found to have a right ear advantage (REA) in the dichotic listening task but left-handers had relatively smaller left/right differences in all of the performance measures. Sample characteristics suggest that there are more IN male right-handers than IN female right-handers.     

Perceptual asymmetries in left- and right-handers for cartoon and real faces

We examined 40 left-handers and 40 right-handers on two free-vision tests of face processing. A chimeric face composed of a smiling half-face joined to either a neutral half-face (real faces) or a sad half-face (cartoon faces) and its mirror image were presented on each trial. Subjects judged which chimeric face looked happier, the one with the smile to the left or the one with the smile to the right. Right-handers, but not left-handers, had a highly significant leftward attentional bias, since chimeras with the smile to the left were judged happier than those with the smile to the right. The cartoon- and real-face tasks did not differ in the mean perceptual asymmetries they elicited. However, correlations between attentional-asymmetry scores for the two tasks, although high for both left- and right-handers, were significantly smaller than task reliabilities. Thus, the cartoon- and real-face tasks overlap to a major extent in the lateralized processes they measure, but to a lesser extent, they also index different lateralized processes.     

Mental rotation and handedness: differences in object-based and egocentric transformations

This study investigates mental rotation performance of right- and left-handers in object-based and egocentric mental rotation tasks using human body stimuli with an outstretched arm in front and back view. Previous literature suggests that right-handers show a slightly better mental rotation performance than left-handers. 42 participants, 14 left-handers and 28 right-handers, completed a mental rotation task with object-based and egocentric transformation of a human figure which was displayed either in front or back view. The main result was a three-way interaction between the factors “kind of transformation”, “handedness” and “view” in a way, that right-handers show significantly faster reaction times then left-handers in front view object-based transformations because of the additional in-depth rotation for front view stimuli. This difference disappeared in egocentric tasks due to the modification of onés own perspective to solve the task. The results of this study show that right-handers not generally outperform left-handers in mental rotation tasks but only if more cognitive resources are needed.     

Hemispheric competition in left-handers on bimanual reaction time tasks

The authors examined possible differences in left- and right-handers on bimanual reaction times to centralized visual stimuli. Eighty participants (n = 40 in each group of left- and right-handers) were tested on unimanual and bimanual reaction time (RT) tasks. Consistently across the 2 groups, the dominant-hand RT was faster, on average, than the nondominant-hand RT, and unimanual RTs were faster than bimanual RTs. However, RT differences between hands revealed a higher percentage of dominant-hand-led trials in right-handers than in left-handers, despite similar absolute RT differences in the 2 groups. On the basis of those findings, the authors conclude that hand dominance does not generally determine which hand leads in a bimanual task and that left-handers have stronger between-hemisphere competition than right-handers do.     

Effects of Fencing Training on Motor Performance and Asymmetry Vary With Handedness

Abstract

Previous studies showed that motor asymmetries are reduced in left-handers and after a long-term fencing training in right-handers. Interestingly, left-handed athletes are substantially over-represented in elite fencing. These findings have been speculatively explained by imbalance in experience of fighting opposite handedness opponents resulted from skewed distribution of handedness, i.e. lefties encounter more righties than righties encounter lefties. Whereas these assumptions could be accurate, the underlying mechanisms remain ambiguous. In this study, we investigated effects of fencing training on motor performance and asymmetry with respect to handedness. We compared fencing performance of left- and right-handed fencers in both training and combat conditions. In the combat condition, left-handers won seven out of twelve matches consisted of twelve bouts each. They also showed a significantly longer hit detection time, a measure indicating better quality of fencing attack. In the training condition, left-handed fencers completed fencing board tests significantly faster than right-handers. These findings provide additional factor of superior motor performance to be considered when interpreting over-representation of lefties in elite fencing. Furthermore, our left-handers were less lateralized, which could explain that superior motor performance. This idea is consistent with previous findings of reduced asymmetry in right-handed fencers when comparing to non-athletes.     

Speech dominance and handedness in the normal human

Spectral analysis was used to measure the coherence or similarity of form between occipital and temporal evoked potentials. In both right- and left-handers, coherence was greater in the left hemisphere for click stimuli and in the right hemisphere for flash stimuli. Similar asymmetries occurred in left but not right speech dominant patients whose speech lateralization has been determined by the intracarotid amytal test. Right-handers and males, however, showed significantly larger amplitudes of auditory responses in the right hemisphere. Left-handers and females reversed this pattern. It was concluded that within a basically left speech dominant organization, males and right-handers would emphasize the verbal, temporal structure of auditory information and the nonverbal, spatial structure of visual information. Females and left-handers would tend to reverse this emphasis.     

Performance Asymmetries in Computer Mouse Control of Right-Handers, and Left-Handers with Left- and Right-Handed Mouse Experience

Precision and general computer mouse aiming performance by right-handers (RH-RM) and left-handers with right-handed mouse experience (LH-RM) and by left-handers with left-handed mouse experience (LH-LM) were compared. A number of performance measures, such as reaction time, time to reach a target, time to click on target, and cursor trajectory, were analyzed. Superficially, specific hand experience seemed to dictate performance asymmetries, but a closer look revealed interactions between hand preference and hand performance. That finding has implications for theories of handedness. In addition, precision and general directional aiming with the mouse cursor showed a clear right hand superiority in reaction time in both RH-RM and LH-RM subjects, whereas LH-LM subjects showed no lateral asymmetries. Finally, the overall time taken for the task, averaged across groups and conditions, favored the experienced hand by some 180 ms. In practical terms, that is not a large difference, especially because the difference will be reduced with practice. Thus, the use of the inexperienced hand can be advocated when there is a need to forestall or ameliorate repetitive stress in the experienced hand.     

Asymmetries in processing horizontal and vertical dimensions

The aim of this study was to determine whether the referential term on the horizontal dimension corresponds to the dominant hand and whether the referential term on the vertical dimension is independent of handedness. In order to verify the hypothesis, right-handers, left-handers, and ambidextrous subjects were required to verify and falsify statements including the words ABOVE and BELOW and the words LEFT and RIGHT. The results showed that right-handers were faster in verifying and falsifying the statements containing the term RIGHT, whereas left handers were faster in verifying and falsifying those containing the term LEFT. Ambidextrous subjects, however, showed no sign of asymmetry in the positional judgments of stimuli along the horizontal dimension. By contrast, right-handers, left-handers, and ambidextrous subjects were equally faster in verifying and falsifying the statements containing the term ABOVE than the statements containing the term BELOW. The relation between the positive term on the horizontal dimension and the dominant hand can be explained by the fact that, in the absence of any asymmetries in the physical world, the dominant hand can furnish a natural reference direction for judgments related to this dimension.     

Crossed-uncrossed difference (CUD) on reaction time and movement time in three two-choice experiments

The present study addresses the robustness and the reliability of the crossed-uncrossed difference (CUD) on a reaction time (RT) and a movement time (MT) component of a prescribed unimanual response to lateralized stimuli. Experiment 1 demonstrated positive CUDs both when a visual warning signal (WS) and an auditory reaction signal (RS) appeared on corresponding and non-corresponding sides of the body. Experiment 2 showed effect of handedness on CUD calculated among right-handers and left-handers. Experiment 3 investigated CUDs through five successive days, indicating that CUDs became steady and reliable although practice affected both RT and MT. All experiments indicated CUD on RT and MT. In addition, Experiments 1 and 2 demonstrated hemispheric asymmetries in favour of an anatomical theory while Experiment 3 did not show any asymmetries and supports an S-R compatibility theory.     

Attentional distribution of task paramters to the two hands during bimanual performance of right- and left-handers

The author tested 12 left-handers and 12 right-handers on a bimanual circling task to examine how attention (either visual or nonvisual) to the task of 1 hand affects within-hand task parameters and whether the effects of attention manipulations are similar in left- and right-handers. The novel prediction that the attended task would be produced larger than the unattended task was confirmed in both handedness groups. The magnitude of the effect on circle size was more pronounced under visual than under nonvisual attention manipulations. The primary effects of attention were similar in the 2 handedness groups, although left-handers demonstrated some evidence of stronger parameter coupling between hands than right-handers did.     

Mirror-reading in right- and left-handers

In mirror-reading, words are read from right to left and letters are read in a reverse right-to-left orientation. In one experiment we compared the ability of normal right- and left-handed subjects to mirror-read and found that the left-handers made fewer errors and could read mirror print more rapidly. In a second experiment we attempted to learn whether there is a hemifield superiority for reading mirror words and whether there are any differences between left- and right-handers in a hemifield. We found that although both right- and left-handers more rapidly detected mirror words projected to the left visual half-field, there were no differences between groups. However, in the right visual half-field, the performance of left-handers was superior to that of the right-handers. The results of the hemifield study suggest that left-handers may be superior at reading mirror words because they can more easily reverse their scanning pattern.     

Lateralization of bilateral transfer of visuomotor information in right-handers and left-handers

The author examined the lateralization of transfer of visuomotor information between the right and left hands during unimanual finger-tapping sequences with visual feedback. The finger-tapping task consisted of a target peak force of 2 N and a target intertap interval of 500 ms. Twenty right-handed and 10 left-handed participants performed the motor task, with 3 transfer trials following 3 practice trials. The author observed positive transfers from the left to the right hand for right-handers but the opposite direction of positive transfers for left-handers. However, left-handers showed a less variable peak force than right-handers did. The author discusses left-handers' interhemispheric information processing.     

Cerebral control for speech in right-handers and left-handers: an analysis of the views of Paul Broca, his contemporaries, and his successors

According to several recent historical accounts, Broca (1865a) stated that left-handers are the mirror-reverse of right-handers for cerebral control of speech, with the right hemisphere being dominant in left-handers, and the left hemisphere dominant in right-handers. The same accounts then note Broca's error in light of current evidence that the majority of left-handers are left-dominant for speech just as are nearly all right-handers. Eling (1984) has called such statements misrepresentations of Broca's position and has argued that Broca's analysis actually was more compatible with the current view that there is a disjunction, meaning an absence of an intimate anatomical relationship, between cerebral control for handedness and speech. The current paper looks again at Broca's work, describes the context in which his views were first articulated, and traces the development of the mirror-reversal principle. The conclusion is reached that, judged by a narrow reading of the 1865 paper, Broca's views could indeed be construed as an anticipation of the modern disjunction principle. However, judged by a broader reading, by consideration of his other writing, and in the context of the philosophical and scientific tradition that shaped his work, it is suggested that it was the mirror-reversal principle to which Broca was actually disposed.     

Left-handedness and achievements in foreign language studies

Studies of brain lateralization lend support to the hypothesis that language-motor functions in left-handers are differently organized from those in right-handers. However, the implications of these differences regarding cognitive functioning are as yet subject to controversy. This concerns all hypotheses raised and empirical data collected over the years. Although it was suggested that left-handers are at higher risk of having language and reading deficits, empirical data from clinical and nonclinical populations are inconclusive at the present time. No effort, however, has been invested in examining possible differences in academic studies of foreign languages according to handedness. Here we report data indicating inferior achievements of left-handed native Hebrew speakers in studies of English as a foreign language. Left-handed pupils significantly more than right-handers were placed in lower level English classes and had more difficulties in applying orthographic-phonological mapping rules in reading English words and pseudowords. However, left-handers' difficulties in this task were not correlated with their performance in a word recognition task. It is thus suggested that the "common symptom" of poor word reading in left-handers indicates different processing failures in different left-handers, some of which impede the buildup of an internal representational system of mapping orthography to phonology and some of which concern mainly the precision of word production.