Human Handedness Research Articles

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.

Hand preference and hand skill in families with schizophrenia

Direction and degree of handedness in humans are variable between individuals and thought to be in part inherited. Several studies have shown an increase in non-right handedness among patients with schizophrenia, and some have included unaffected relatives. The present study was designed to determine whether reduced right handedness is more frequent among individuals with schizophrenia as compared with their well relatives and whether it clusters within families having multiple ill members. A total of 259 families comprising 418 individuals diagnosed with schizophrenia or schizoaffective disorder, 54 individuals with other psychoses, 145 family members with depression and other minor diagnoses, and 288 unaffected individuals were included. Hand preference was assessed by the Annett Scale and right relative to left hand skill measured using the Tapley-Bryden test. For all assessments of hand preference and hand skill, females were significantly more lateralised towards the right than males. Those individuals with schizophrenia or schizoaffective disorder had significantly less right hand preference than their unaffected relatives when measured as a quantitative index of items from the Annett Scale (p = .019), but not categorically (right, left or mixed). In contrast, there was no difference in hand skill between diagnostic groups. Hand preference was significantly correlated among male-male affected sibling pairs (p = .01) and similar results were found for hand skill among the total group of affected pairs (p = .001). Although these results only partially support a relationship between handedness and schizophrenia, they nevertheless draw attention to sex differences in hand preference and the familial aspects of hand preference in this disorder. More direct approaches to the genetics of cerebral dominance and psychosis are required.

Paw preference in rats.

We applied a food-reaching test to investigate paw preference in 114 Spraque-Dawley rats and to determine whether a correlation exists between paw preference and gender. We found that 70.2% of rats were right-pawed, 19.3% were left-pawed, and 11.9% were ambidextrous. No significant difference was found between right-pawed, left-pawed, and ambidextrous male and female rats. The results indicate that the distribution of paw preference in rats is similar to that of other animals and to human handedness.

Inferring handedness from lithic evidence

Until recently research into the origins of human handedness has been hampered by the lack of valid techniques for inferring handedness in pre-modern populations. A method developed by Toth for inferring handedness from lithic evidence, based on orientation of the cortex on lithic flakes, has produced promising results. However, this method is limited in applicability and has a variable signal to noise ratio. The authors describe a separate method, based on the orientation of the cone of percussion in lithic flakes, for inferring handedness from the lithic evidence. This method complements the cortex method. Some preliminary experimental evidence is presented which indicates that handedness can be inferred from lithic evidence using the cone of percussion method. Suggestions for further research are made.

Postnatal development of lateralized motor preference in the African Grey parrot ( Psittacus erithacus)

The parrot appears to provide a potentially unique animal model of handedness in humans, but few (if any) observational studies of early postnatal development of postural/motor asymmetries have been published. We studied three African Grey hatchlings, raised without human physical contact, for the first 5 months of life. All three animals failed to show consistent postural and/or motor asymmetries until the end of the 4 postnatal week. These results appear to be comparable to data from prior studies with human infants. Delayed development of lateral motor and/or postural preferences may represent an evolutionarily adaptive strategy for altricial animals.

Genes for left-handedness: How to search for the needle in the haystack?

Although several genes that determine left-right asymmetry for structural syndromes such as situs inversus have been characterised in recent years (Supp, Witte, Potter, & Brueckner, 1997), there has been little progress in determining which genes or loci predispose to left-right handedness in humans. Linkage analysis has been used widely for the localisation of genes followed by their positional cloning. The complex genetics of handedness is one of the greatest problems for standard linkage analysis. Several genetic models have been proposed for the inheritance of handedness in humans. On the basis of these models, lefthandedness can be considered a common single gene trait with a high gene frequency and a non-mendelian inheritance pattern. We report here a possible strategy, using these genetic models, that can be applied for the identification for genes determining handedness in humans.

Birth order and left-handedness revisited: some recent findings in chimpanzees ( Pan troglodytes) and their implications for developmental and evolutionary models of human handedness

This paper examines the relationship between parity, pregnancy outcome, and handedness in a sample of captive chimpanzees ( Pan troglodytes). The relation between parity, maternal age and positive or negative pregnancy outcome was assessed from life history data for 536 chimpanzees housed at the Yerkes Regional Primate Research Center. The incidences of negative pregnancy outcome (notably spontaneous abortions and stillbirths) were significantly higher in parities of 8 or higher compared to all other parities. In a sub-sample of 165 chimpanzees, the relation between parity, maternal age and handedness was assessed to determine whether left handedness may serve as a marker of prenatal pathology. These analyses indicated that left-handedness was more prevalent in 1st and 8 or higher parities compared to parities between 2 and 7, respectively. Possible prenatal hormonal and periparturitional factors are discussed as possible mechanisms for the observed findings.

Handedness in past human populations: Skeletal markers

Although it is generally believed that right-handedness has predominated in all human populations since at least the appearance of Homo sapiens sapiens, the evidence has rarely been studied in its entirety. This paper contains a review of the principal skeletal indicators of handedness (cranial and post-cranial), and an introduction to their use to estimate the prevalence of left and right handedness in the past. The role of such reviews is to enable us to assess the testability of genetic models of human handedness and of its evolution, using anthropological and archaeological data.

On the Evolution of Handedness: A Speculative Analysis of Darwin's Views and a Review of Early Studies of Handedness in “the Nearest Allies of Man”

Scientists today who seek clues into the evolutionary origins of human handedness make extensive use of evidence from comparative studies, that is, studies that ask whether handedness occurs in other species, especially apes and monkeys, as the Darwinian principle of continuity would seem to imply, or whether it is uniquely human. Early investigations had the same goal and drew on much the same kind of evidence. In this article, I review studies of animal handedness in the period before 1859, when Darwin published On the Origin of Species, and afterward, through the 1st decade of the 20th century. Inasmuch as Darwin's published writings contain hardly any statements about handedness and none at all about its evolution and continuity across species, I also speculate about what Darwin himself might have said on the subject. To do this, I draw on his statements on related matters, such as the form and structure of the hand and the transition from a quadrupedal to bipedal stance, on other writers' reports and opinions about handedness with which he was familiar or likely to have been familiar, and finally, on clues from his own and only statement about animal handedness in an unpublished letter. I conclude by asking whether and how early investigators, lacking any statement by Darwin on the evolution of handedness, invoked his theory of evolution and his views on related matters in the interpretation of their findings.

Foundations of a Functional Theory of Human Handedness

A functional theory of human handedness is introduced based on early childhood experiences and the neural plasticity of the brain. It assumes that very different factors may be involved in the development of an individual left-handedness. Imitation and social learning are major factors in the formation of both right- and left-handedness in normal early childhood development. However, the frequently claimed relation between left-handedness and pathology as well as exceptional giftedness can only be understood by taking into account the function or meaning of left-handedness for personality development. The functional approach conceives left-handedness as a ubiquitous symbolic aid for coping creatively with conflict in early childhood. A childrearing style that clashes with the child's body-mind needs can lead the socioculturally less favored left hand to serve as a symbolic frame of reference for a meaning-making action repertoire that is fundamental in protecting and stabilizing the childhood self and personality development. Suggestions are derived regarding brain lateralization and handedness and for the diagnosis of early childhood disorders.

Handedness and aggressive behavior in an Ntumu village in southern Cameroon

Handedness in humans is possibly maintained by a frequency-dependent advantage during aggressive interactions. To contribute to relevant data related to this idea, functional handedness has been measured in a traditional ethnic group (Ntumu) from southern Cameroon, and confronted to the level of aggressive behavior. Functional handedness was measured by the hand holding the machete, an asymmetric tool used every day. Aggressive behavior was assessed by the frequency of occurrence of the traditional tribunal for solving internal conflicts. There was a low percentage of left-handers (8.1%), with no significant sex differences, although females below 50 year old displayed a significant increase of left-handedness with age. The level of aggressive behavior was low, as the traditional tribunal took place only seven times during the last five years. Results are discussed in the context of the evolution of handedness and the possible recent societal changes following European colonisation.

Birth order and hand preference in chimpanzees (Pan troglodytes): implications for pathological models of handedness in humans.

The effect of birth order on hand preference was assessed in a sample of 154 captive-born chimpanzees. Subjects were classified as first, middle, or latter born using 2 classification criteria based on their birth order. Hand preference was measured using a task that elicited coordinated bimanual actions. Significant birth-order effects were found for both classification criteria, with first- and latter-born subjects exhibiting a lesser degree of right-handedness compared with middle-born subjects. These data suggest that biological rather than sociological factors play a greater role in explaining the observed birth-order effects on hand preference in humans.

Birth order and hand preference in chimpanzees (Pan troglodytes): Implications for pathological models of handedness in humans.

Birth order and hand preference in chimpanzees (Pan troglodytes): Implications for pathological models of handedness in humans.

Human handedness and asymmetry of the motor cortical silent period.

We performed transcranial magnetic stimulation of the motor cortex in 22 left-handed and 25 right-handed subjects during active contraction of a small hand muscle. Motor evoked potentials had the same latency, amplitude and threshold on both sides of the body, whilst the silent period duration was shorter in the dominant hand. Silent periods elicited by nerve and brainstem stimulation were the same in both hands. Since the latter part of the cortical silent period is due mainly to withdrawal of corticospinal input to spinal motoneurones, we speculate that the results are compatible with the suggestion that tonic contractions of the non-dominant hand are associated with a greater involvement of the corticospinal tract than those of the dominant hand. It also seems likely that there is an asymmetry in the excitability of cortical inhibitory mechanisms with those responsible for the cortical silent period being less excitable in the dominant motor cortex.

Hand preference and transcranial magnetic stimulation asymmetry of cortical motor representation

Human handedness may be associated with asymmetry in the corticospinal motor system. Previous studies measuring the threshold for eliciting motor evoked potentials (MEPs) to transcranial magnetic stimulation (TMS) have provided evidence consistent with this hypothesis. However, TMS asymmetry observed in previous studies may have reflected cortical or spinal differences. We therefore undertook this investigation to test the hypothesis that handedness is associated with asymmetry in cortical motor representations. We used TMS to map contralateral cortical motor representations of the right and left abductor pollicis brevis (APB) and flexor carpi radialis (FCR) muscles in nine normal subjects (three left-handed). Using focal stimulation with a figure-of-8 shaped magnetic coil, we found no differences in MEP threshold or MEP size between the preferred and the nonpreferred hand. However, we observed that the number of scalp stimulation sites eliciting MEPs was statistically greater for APB and FCR muscles of the preferred limb. We found significant asymmetry between right-handed and left-handed subjects, such that in right-handers, the representation of the right APB was larger than that of the left APB, but in left-handers the representation of right APB was smaller than that of the left APB. These results suggest that handedness is associated with asymmetry in cortical motor representation.

Handedness-related asymmetry in transmission in a system of human cervical premotoneurones.

The possibility was investigated that human handedness is associated with an asymmetrical cortical and/or peripheral control of the cervical premotoneurones (PreMNs) that have been shown to mediate part of the descending command to motoneurones of forearm muscles. Heteronymous facilitation evoked in the ongoing voluntary extensor carpi radialis (ECR) electromyographic activity (EMG) by weak (0.8 times motor threshold) stimulation of the musculo-cutaneous (MC) nerve was assessed during tonic co-contraction of biceps and ECR. Suppression evoked by stimulation of a cutaneous nerve (superficial radial, SR) at 4 times perception threshold in both the voluntary EMG and in the motor evoked potential (MEP) elicited in ECR by transcranial magnetic stimulation (TMS) was investigated during isolated ECR contraction. Measurements were performed within time windows or at interstimulus intervals where peripheral and cortical inputs may interact at the level of PreMNs. Results obtained on both sides were compared in consistent right- and left-handers. MC-induced facilitation of the voluntary ECR EMG was significantly larger on the preferred side, whereas there was no asymmetry in the SR-evoked depression of the ongoing ECR EMG. In addition, the suppression of the ECR MEP by the same SR stimulation was more pronounced on the dominant side during unilateral, but not during bilateral, ECR contraction. It is argued that (1) asymmetry in MC-induced facilitation of the voluntary EMG reflects a greater efficiency of the peripheral heteronymous volley in facilitating PreMNs on the dominant side; (2) asymmetry in SR-induced suppression of the MEP during unilateral ECR contraction, which is not paralleled by a similar asymmetry of voluntary EMG suppression, reflects a higher excitability of cortical neurones controlling inhibitory spinal pathways to cervical PreMNs on the preferred side.

Human handedness: an ethological perspective

Human handedness is thought to be unique and universal. Critical examination of primary sources challenges both of these conclusions. Data on laterality of function in apes, humankind’s closest living relations, indicates hand preference and task specialization, but not yet handedness, as crucial studies remain to be done. Ethnographic data across cultures reveal more variance than is usually acknowledged, but existing findings are based on minimal and limited data. Ethological studies of spontaneous behavior in every day life are needed for both human and non-human primates.

On the Genetics and Measurement of Human Handedness

Lateral preferences in humans and other primates may be a key to many important issues in biology and psychology. There is strong evidence that the representation of language functions in the cerebral cortex is different in left-handed (LH) people than in right-handed (RH) people, and an understanding of handedness may lead to valuable clues as to how the brain becomes organised in the way that it does. Although there are clearly cultural effects that influence manual activities, there is nevertheless evidence that human handedness and other lateralities are at least in part genetically determined: compared with two RH parents, one RH and one LH parent are 2-3 times more likely to have an LH child, and two LH parents are 3-4 times more likely to have an LH child. Thus, one might wish to investigate the genetics of laterality with the goal of understanding the biological mechanisms that lead to the preferential use of one hand (or eye or foot). One may also see links between human handedness and footedness and the motor asymmetries found in many nonhuman primate species, and perhaps in lower mammals as well. From this perspective, one might see the study of human laterality as relevant to evolutionary biology. Investigators of both human and nonhuman asymmetries have to grapple with such difficult measurement issues as the relations between preference and performance, and the influences of postural and task demands, and have much to offer each other in the quest for the nature of laterality. Our recent work seems to indicate that the various sensory and motor lateralities may be related, in humans, but not in a simple way. In future work, the challenge will be to identify the relations between the various laterality ''profiles'' and patterns of functional cerebral organisation.

On the Genetics and Measurement of Human Handedness

On the Genetics and Measurement of Human Handedness

The mixture distribution of left minus right hand skill in men and women

The distribution of left minus right (L—R) hand skill (peg moving) was studied in 436 men and 247 women. In total sample, the distribution was continuous and a sum of two Gaussian curves. One of them was constituted by subjects with better right hand (79.0%) and better left hand (21.0%). The other curve (2.5% of total) was exclusively due to the subjects with better left hand. In men and women, the total curve was a sum of three overlapping Gaussian curves. Two of them were mostly due to the subjects with better right hand. There was a third curve representing almost entirely the subjects with better left hand. Women were more strongly right-handed than men. There was no significant sex difference at the sinistral side. The overall results were compatible with the right shift theory, but there was not a single Gaussian curve representing a chance distribution with a mean of zero. These results suggested that the distribution of manual asymmetry in skill is continuous, which may be described by probabilities due to subjects with better right and left hands; there is no place for chance in human handedness.