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Birth order, maternal age, season of birth, and handedness.

The handedness of 1,186 children about to enter kindergarten was tested. The sample represented approximately 98 percent of all children of this age in a school system during 8 years. Handedness was examined in relation to birth order, maternal age at birth, and sex, for 1,097 subjects (92.5%), and in relation to season of birth for the total sample. The finding that there were significantly more nonright-handed boys for birth orders one and four-or-later, than for two and three, provides support for Bakan (1971). There was a nonsignificant increase in nonright-handedness for boys born to the youngest (less than 20) and oldest (less than 30) mothers. A significant seasonal effect for the birth of nonright-handed boys was observed. For each of the fall and winter months (September - February) the proportion of nonright-handed male births was higher than that for any of the spring and summer months. For girls, no significant effects on handedness were observed, for birth order, maternal age, or season of birth.

Birth Order↗

Factoring handedness data: II. Geschwind's multidimensional hypothesis.

The challenge in this journal by Peters and Murphy to the validity of two published factor analyses of handedness data because of bimodality was dealt with in Part I by identifying measures to normalize the handedness item distributions. A new survey using Oldfield's questionnaire format had 38 bell-shaped (unimodal) handedness-item distributions and 11 that were only marginally bimodal out of the 55 items used in Geschwind's 1986 study. Yet they were still non-normal and the factor analysis was unsatisfactory; bimodality is not the only problem. By choosing a transformation for each item that was optimal as assessed by D'Agostino's K2 statistic, all but two items could be normalized. Seven factors were derived that showed high congruence between maximum likelihood and principal components extractions before and after varimax rotation. Geschwind's assertion that handedness is not unidimensional is therefore supported.

Analysis of Variance↗

Neurocognitive impairments associated with ambiguous handedness in the chronically mentally ill.

One form of atypical handedness, ambiguous handedness, is found in roughly one-quarter of chronic schizophrenic patients. Despite its prevalence, relatively little is known about the neurocognitive underpinnings of ambiguous handedness. In the present study we examined the performance of ambiguous (n = 19) and non-ambiguous (n = 39) handed chronically mentally ill inpatients on selected measures of verbal learning, motor learning and manual dexterity. The results revealed that ambiguous handers were more impaired than non-ambiguous handers in verbal learning, but not motor learning. Group differences in manual dexterity were significant for the entire sample, but not when analyses were limited to males. These findings suggest that impairments in verbal learning may be linked to the pathogenesis of ambiguous handedness in chronic psychiatric patients.

Adult↗

Handedness and immune function.

Geschwind, Galaburda, and Behan (GBG) have suggested that in utero levels of testosterone influence both cerebral and immune system developments (Geschwind and Behan, 1982; Geschwind and Galaburda, 1984; Geschwind and Galaburda, 1985). According to this theory, high levels of testosterone result in greater incidences of left-handedness, deviations from standard distribution of cerebral functions (known as anomalous dominance), and increased autoimmune dysfunction. While the original data supported these assertions, more recent tests of the hypothesis have been equivocal. One criticism of these studies is that the definition of both handedness and anomalous dominance are too vague. It was one of the aims of this project to investigate and clarify the GBG model by examining four different aspects of handedness as well as a more direct measure of anomalous dominance. In order to extend the GBG model, degree of left-handedness, general immune system functioning, and current testosterone levels were also examined. First, it was predicted and found that left handers had a higher incidence of autoimmune diseases in their immediate families than did right handers. Second, those left handers with an incidence of at least one autoimmune disease were more strongly left-handed than were those with no incidence of autoimmunity. Finally, it was observed that higher testos terone levels were supportive of general immunity. The present findings both support and expand the GBG model.

Adult↗

Routine ultrasound screening in pregnancy and the children's subsequent handedness.

OBJECTIVE: To study a possible association between ultrasound screening in early pregnancy and altered cerebral dominance measured by the prevalence of non-right handedness among children, particularly boys. METHODS: Follow-up of 8 to 9 year old children to women who participated in a randomised controlled trial on ultrasound screening during pregnancy in 1985-87. The children were followed up through a questionnaire sent to their mothers. The dominant hand of the child was assessed by eleven questions. The dominant foot by one question. RESULTS: No differences were found in non-right handedness between children in the screening and non-screening group. In separate analyses on ultrasound exposure and non-right handedness among boys a significant difference was found (odds ratio 1.33; 95% confidence interval 1.02-1.74). CONCLUSION: This study could not rule out a possible association between non-right handedness among boys and ultrasound exposure in early fetal life. The association was, however, confined to analyses comparing exposed and non-exposed boys and no associations were found when the comparisons were performed according to the randomised groups.

Child↗

Handedness and corpus callosum morphology.

Investigations of a relationship between callosal size and functional behavioral lateralization lead to the hypothesis that, as the size of the corpus callosum (CC) increases, interhemispheric information transfer is facilitated and behavioral laterality effects become smaller. The aim of our in vivo study was to investigate the relationship between functional asymmetry of handedness and CC size in healthy subjects. Magnetic resonance images of the CC and five CC subregions were obtained with a 1.5-T Magnetom using a three-dimensional T1 sequence in 46 healthy men. Handedness was determined using the 'handedness dominance test' (HDT). According to the HDT values, 32 consistent and 14 non-consistent right-handers were identified. No significant difference between handedness subgroups in CC regions and no significant correlations between HDT values and CC areas were detected.

Adolescent↗

The relationship of age, gender, handedness, and sidedness to the size of the corpus callosum.

OBJECTIVES: We correlated the area and size of the corpus callosum, as measured by MR imaging, with the individual's handedness, sidedness, age, and gender. MATERIALS AND METHODS: A total of 117 patients (59 male, 58 female) aged 15-75 years were selected for this study. This included 12 persons who were either left-handed or ambidextrous. Each patient was tested extensively to determine handedness and sidedness. Callosal areas and thickness were measured and correlated with brain size. RESULTS: The body of the corpus callosum decreases in size with age and is larger in right-handed persons. The cross-sectional areas of the genu, splenium, and corpus callosum, overall, do not vary significantly with respect to age, gender, sidedness, or handedness. CONCLUSIONS: The size of the corpus callosum consistently decreases with age. Otherwise, few statistical differences in callosal size relate to gender, sidedness, or handedness.

Adolescent↗

Ability of viral topoisomerase II to discern the handedness of supercoiled DNA: bimodal recognition of DNA geometry by type II enzymes.

Previous studies with human and bacterial topoisomerases suggest that the type II enzyme utilizes two distinct mechanisms to recognize the handedness of DNA supercoils. It has been proposed that the ability of some type II enzymes, such as human topoisomerase IIalpha and Escherichia coli topoisomerase IV, to distinguish supercoil geometry during DNA relaxation is mediated by elements in the variable C-terminal domain of the protein. In contrast, the ability of human topoisomerase IIalpha and topoisomerase IIbeta to discern the handedness of supercoils during DNA cleavage suggests that residues in the conserved N-terminal or central domain of the protein are involved in this process. To test this hypothesis, the ability of Paramecium bursaria chlorella virus-1 (PBCV-1) and chlorella virus Marburg-1 (CVM-1) topoisomerase II to relax and cleave negatively and positively supercoiled plasmids was assessed. These enzymes display a high degree of sequence identity with the N-terminal and central domains of eukaryotic topoisomerase II but naturally lack the C-terminal domain. While PBCV-1 and CVM-1 topoisomerase II relaxed under- and overwound substrates at similar rates, they were able to discern the handedness of supercoils during the cleavage reaction and preferentially cut negatively supercoiled DNA. Preferential cleavage was not due to a change in site specificity, DNA binding, or religation. These findings are consistent with a bimodal recognition of DNA geometry in which topoisomerase II uses elements in the C-terminal domain to sense the handedness of supercoils during DNA relaxation and elements in the conserved N-terminal or central domain during DNA cleavage.

DNA Topoisomerases, Type II↗

Variations in cerebral organization as a function of handedness, hand posture in writing, and sex.

During the past century, it has become increasingly apparent that there is a great deal of variation in the direction and degree of cerebral lateralization, a plurality of people having language and related functions strongly specialized to the left hemisphere and visuospatial functions strongly specialized to the right, with substantial minorities manifesting various deviations from this pattern. In particular, in 35%-50% of sinistrals and 1%-10% of dextrals, the right hemisphere is specialized for linguistic skills, and in some unknown fraction of the two handedness groups, verbal and/or spatial abilities are, to varying extents, bilateralized. Levy (1973) suggested that the hand posture adopted during writing might be an index of the lateral relationship between the dominant writing hand and the language hemisphere, a normal posture indicating contralateral language specialization, and an inverted posture indicating ipsilateral language specialization. In the present investigation, two tachistoscopic tests of cerebral lateralization, one measuring spatial functions and one measuring verbal function, were administered to 73 subjects classified by handedness, hand posture during writing, and sex. Among both dextral and sinistral subjects with a normal writing posture, language and spatial functions were specialized to the contralateral and ipsilateral hemispheres, respectively, and lateral differentiation of the brain was strong. The reverse was seen in subjects having an inverted writing posture. In all groups, females were less laterally differentiated than males. In 70 out of 73 subjects, the direction of cerebral laterization was accurately predicted by handedness and hand posture. The 3 subjects (2 females and 1 male) who failed to manifest the predicted relations were all left-handers having an inverted hand posture . In this group, lateral differentiation was so weak that the reliability of the tachistoscopic tests was reduced, and we attribute these three predictive failures to this cause. Thus, almost all of the variation in the lateral organization of the brain was accounted for by handedness, hand posture, and sex.

Cerebral Cortex↗

Evidence for the pathological right-handedness hypothesis.

The present study investigated handedness of 109 Korean patients with temporal lobe epilepsy (TLE) and 120 normal control participants. Left TLE was associated with a significantly raised incidence of left-handers relative to normal controls. More important for the present purpose, right TLE was associated with a significantly raised incidence of strong right-handers relative to normal controls. This finding indicates that certain early damage to the right brain causes a mild hypofunction of the left hand in natural weak right-handers, which in turn causes the patient to switch to strong right-handedness. To the authors' knowledge, this finding provides the 1st empirical evidence supporting the pathological right-handedness hypothesis. Prior failure to find evidence of pathological right-handedness may reflect the low base rate of natural left-handers rather than true absence of pathological right-handers.

Adult↗

Chimpanzee (Pan troglodytes) handedness: variability across multiple measures of hand use.

This study examined intertask consistency in handedness across multiple measures of hand use in a sample of 187 chimpanzees (Pan troglodytes). Hand preferences for 2 to 6 measures were collected from the sample, and hand preference scores were derived on the basis of the individual hand preferences for each measure. Seven of 15 possible intratask correlations were significant, with some degree of clustering depending on the motor demands of the tasks. Two overall measures of handedness revealed population-level right-handedness in the chimpanzees, although the degree of bias was reduced for chimpanzees tested on more than 3 measures of hand use. The results are interpreted in the context of several recent studies that proposed theoretical models of handedness in nonhuman primates.

Analysis of Variance↗

An unconventional myosin in Drosophila reverses the default handedness in visceral organs.

The internal organs of animals often have left-right asymmetry. Although the formation of the anterior-posterior and dorsal-ventral axes in Drosophila is well understood, left-right asymmetry has not been extensively studied. Here we find that the handedness of the embryonic gut and the adult gut and testes is reversed (not randomized) in viable and fertile homozygous Myo31DF mutants. Myo31DF encodes an unconventional myosin, Drosophila MyoIA (also referred to as MyoID in mammals; refs 3, 4), and is the first actin-based motor protein to be implicated in left-right patterning. We find that Myo31DF is required in the hindgut epithelium for normal embryonic handedness. Disruption of actin filaments in the hindgut epithelium randomizes the handedness of the embryonic gut, suggesting that Myo31DF function requires the actin cytoskeleton. Consistent with this, we find that Myo31DF colocalizes with the cytoskeleton. Overexpression of Myo61F, another myosin I (ref. 4), reverses the handedness of the embryonic gut, and its knockdown also causes a left-right patterning defect. These two unconventional myosin I proteins may have antagonistic functions in left-right patterning. We suggest that the actin cytoskeleton and myosin I proteins may be crucial for generating left-right asymmetry in invertebrates.

Actins↗

Parametric and non-parametric linkage analysis of several candidate regions for genes for human handedness.

The frequency of left-handedness in the general population is around 11%. Both environmental and genetic models have been proposed to explain the aetiology of human handedness. The majority of genetic models, such as those of Annett, McManus and Klar, propose a single gene determinant with a non-Mendelian inheritance pattern. As left-handedness is correlated with cerebral asymmetry and is a feature of left-right asymmetry, genes involved in the development of left-right asymmetry can be considered as candidate genes. Candidate gene analysis was performed using an informative extended pedigree, and also using nuclear families of right-handed parents with left-handed children. Segregation analysis in the extended pedigree identified allele sharing in the NODAL and DNAHC13 candidate regions on chromosome 10 and 1. Linkage analysis using the models of Klar and McManus, and non-parametric analysis on nuclear families, subsequently excluded all candidate regions tested. This demonstrates the power to identify the genes specifying handedness by the conduct of extended genetic studies on these and similar cohorts.

Axonemal Dyneins↗

Condylar asymmetry and handedness in patients with temporomandibular disorders.

The relationship between condylar asymmetry and handedness of dentate patients with temporomandibular disorders was investigated. A group of 28 patients (22 females and 6 males) was studied. The mean age was 28.29 years, with a range from 10 to 54 years. The association between handedness and the sign of the condylar asymmetry index was examined using Fisher's exact test. No difference was found between sign and handedness (P > 0.05). This suggests that handedness does not affect the asymmetry index in this group of patients.

Adolescent↗

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.

Acoustic Stimulation↗

A code-theoretical note on object handedness.

This study is a theoretical exercise dealing with discrimination between images and mirror-images. It focuses on the way codes of shapes represent their handedness. We compare two code systems with different reference frames. These frames determine the specific sensitivity of each system. One system uses an asymmetric reference frame. It is called the H-system and was inspired by an idea of Corballis (1988 Psychological Review 95 115-123). The other system, being our proposal, uses a symmetric reference frame and we have named it the M-system. We demonstrate the following. A code of the H-system provides a cue for the handedness of a shape, but not for rotation, i.e. no cue for the appropriate kind of code rotation which should be tested in case images and mirror-images are discerned by mental rotation. The M-system is the converse in both respects. A code of this system does not provide a handedness cue but, instead, a rotation cue. Thus, for handedness discrimination, the H-system neither needs nor guides mental rotation, whereas the M-system does both. This M-system generates object-centred structural codes enriched with viewpoint information. Various visual experiments reported in the literature favour the M-system over the H-system, implying that perception does not make use of an asymmetric but of a symmetric reference frame.

Form Perception↗

Heritability of lobar brain volumes in twins supports genetic models of cerebral laterality and handedness.

Although the left and right human cerebral hemispheres differ both functionally and anatomically, little is known about the environmental or genetic factors that govern central nervous system asymmetry. Nevertheless, cerebral asymmetry is strongly correlated with handedness, and handedness does have a significant genetic component. To explore the relative contribution of environmental and genetic influences on cerebral asymmetry, we examined the volumes of left and right cerebral cortex in a large cohort of aging identical and fraternal twins and explored their relationship to handedness. Cerebral lobar volumes had a major genetic component, indicating that genes play a large role in changes in brain volume that occur with aging. Shared environment, which likely represents in utero events, had about twice the effect on the left hemisphere as on the right, consistent with less genetic control over the left hemisphere. To test the major genetic models of handedness and cerebral asymmetry, twin pairs were divided into those with two right handers and those with at least one left hander (nonright handers). Genetic factors contributed twice the influence to left and right cerebral hemispheric volumes in right-handed twin pairs, suggesting a large decrement in genetic control of cerebral volumes in the nonright-handed twin pairs. This loss of genetic determination of the left and right cerebral hemispheres in the nonright-handed twin pairs is consistent with models postulating a right-hand/left-hemisphere-biasing genetic influence, a "right-shift" genotype that is lost in nonright handers, resulting in decreased cerebral asymmetry.

Aged↗

Direction of handedness linked to hereditary asymmetry of a sensory system.

Studies on the role of heredity in the transmission of handedness in nonhuman mammals have, so far, led to the isolation of mouse strains that differed in the lateralized versus ambidextrous use of the forepaw in a food-retrieval task (strength of paw preference). Here we report that left versus right use of the forepaw (direction of paw preference) is associated with a genetically expressed structural asymmetry of a sensory system, the whisker-to-barrel pathway. Mice that express whisker pad asymmetry of a direction that corresponds with the asymmetry for which they were bred demonstrate an opposite shift in the distribution of handedness: a right or left dominance of the whisker pad predicts a high proportion of left-handers or right-handers, respectively. Is an altered brain circuit--that is, a consequence of the asymmetry of the whisker pad--associated with a change in the circuitry that governs handedness? Or, alternatively, are there two gene sets responsible for the phenomena that we report--one that causes "whiskeredness" and another that causes handedness?

Animals↗