Handedness Polymorphism Research Articles

Left-handedness is associated with greater fighting success in humans

Left-handedness is a costly, sexually dimorphic trait found at low frequencies in all human populations. How the handedness polymorphism is maintained is unclear. The fighting hypothesis argues that left-handed men have a negative frequency-dependent advantage in violent intrasexual competition giving them a selective advantage. In support of this, many studies have found that left-handed men are overrepresented among modern professional fighters, but studies typically find no difference in fighting success between left and right-handed fighters. We studied over 13,800 professional boxers and mixed martial artists of varying abilities in three of the largest samples to test this hypothesis to date, finding robust evidence that left-handed fighters have greater fighting success. This held for both male and female fighters, and for both percentage of fights won and an objective measure of fighting ability. We replicated previous results showing that left-handed fighters are strongly overrepresented in professional combat sports, but left-handed fighters did not show greater variance in fighting ability, a hypothesis suggested in previous studies. Overall we find strong evidence consistent with the fighting hypothesis.

Handedness heritability in industrialized and nonindustrialized societies.

In modern societies, there is a decreased usage of traditional weapons to settle interpersonal or inter-group disputes compared to usage in traditional societies, possibly affecting the frequency-dependent selection on the handedness polymorphism. Another societal difference is the extensive automation of hard manual labour (including agriculture) in industrialized societies, relaxing the selection for hand specialization. Thus, selection of handedness is likely to differ between traditional and modern societies. As heritability determines the relative speed of evolutionary dynamics, handedness heritability was compared between industrialized and non-industrialized societies. First, individuals were sampled from a non-industrialized area in Indonesia, where violent conflicts are relatively frequent and tribal wars have been prevalent recently. Handedness was recorded directly or indirectly for 11,490 individuals belonging to 650 independent pedigrees, and handedness heritability was estimated using a pedigree-based animal model. Second, estimates of handedness heritability derived from published sources were collected to compare heritability estimates, accounting for various confounding variables. Non-industrialized countries displayed a significantly higher heritability value (h2 = 0.56) than that of industrialized countries (h2 = 0.20). Heritability decreased with time along the twentieth century in industrialized countries, independently of the frequency of left-handedness, and independently of the method used to measure handedness. In conclusion, the data are consistent with a decrease in handedness heritability following the industrialization process and/or the associated decrease in violence using traditional weapons. The difference in heritability between industrialized and non-industrialized countries suggests that selection of handedness is thus likely to differ between traditional and modern societies.

Left-handedness and time pressure in elite interactive ball games.

According to the fighting hypothesis, frequency-dependent selection gives relatively rarer left-handers a competitive edge in duel-like contests and is suggested as one mechanism that ensured the stable maintenance of handedness polymorphism in humans. Overrepresentation of left-handers exclusively in interactive sports seems to support the hypothesis. Here, by referring to data on interactive ball sports, I propose that a left-hander's advantage is linked to the sports' underlying time pressure. The prevalence of left-handers listed in elite rankings increased from low (8.7%) to high (30.39%) time pressure sports and a distinct left-hander overrepresentation was only found in the latter (i.e. baseball, cricket and table tennis). This indicates that relative rarity and the interactive nature of a contest are not sufficient per se to evoke a left-hander advantage. Refining the fighting hypothesis is suggested to facilitate prediction and experimental verification of when and why negative frequency-dependent selection may benefit left-handedness.

The fighting hypothesis as an evolutionary explanation for the handedness polymorphism in humans: where are we?

The ubiquitous and persistent handedness polymorphism in humans requires an evolutionary explanation. It has been suggested that left-handers have a frequency-dependent advantage during a fight, such that this advantage decreases when their frequency increases. Many independent studies are providing data from interactive sports (a specific class of fights), and are very supportive of the fighting hypothesis. The only intercultural study on traditional societies is also consistent with the fighting hypothesis, although it has not yet been replicated. The frequencies of left-handers in the few remaining violent societies are likely to be rapidly decreasing, due to Western colonization (long-range weapons, religion, and money market) dramatically affecting the frequency-dependent selection associated with handedness. Clearly, more data are urgently needed outside the Western influence.

Human handedness in interactive situations: Negative perceptual frequency effects can be reversed!

Left-handed performers seem to enjoy an advantage in interactive sports. Researchers suggest this is predominantly due to the relative scarcity of left-handers compared with right-handers. Such negative frequency-dependent advantages are likely to appear in inefficient game-play behaviour against left-handed opponents such as reduced ability to correctly anticipate left-handers' action intentions. We used a pre–post retention design to test whether such negative frequency-dependent perceptual effects can be reversed via effective training. In a video-based test, 30 handball novices anticipated the shot outcome of temporally occluded handball penalties thrown by right- and left-handed players. Between the pre- and post-tests, participants underwent a perceptual training programme to improve prediction accuracy, followed by an unfilled retention test one week later. Participants were divided into two hand-specific training groups (i.e. only right- or left-handed shots were presented during training) and a mixed group (i.e. both right- and left-handed shots were presented). Our results support the negative frequency-dependent advantage hypothesis, as hand-specific perceptual training led to side-specific improvement of anticipation skills. Similarly, findings provide experimental evidence to support the contention that negatively frequency-dependent selection mechanisms contributed to the maintenance of the handedness polymorphism.

Left-Handedness and Male-Male Competition: Insights from Fighting and Hormonal Data

Male-male competition can shape some behavioral or morphological traits of males. Here we investigate if this competition could play a role in the persistence of the polymorphism of handedness in human populations. A negative frequency-dependent selection mechanism has been hypothesized, based on the fact that left-handed men may benefit from a "surprise" advantage during fighting interactions because they are rare in human populations. This advantage may thereby enhance the probability of survival of left- handed men and/or their reproductive success through an increase in social status. In this study, we first explored the association between hand preference and lifetime fighting behavior in a population of 1,161 French men. No effect of hand preference on the probability of fighting was detected, suggesting that the innate propensity to fight does not differ between left- and right-handers. However, among men who had been involved in at least one fight during their lifetime, left-handers reported significantly more fights than right-handers. To explore the biological basis of this behavior, we also investigated the testosterone concentration in saliva samples from 64 French university students. Consistent with frequencies of fights, we found a significantly higher average testosterone concentration in left-handers than in right-handers. We suggest that these behavioral and hormonal differences may be acquired throughout life due to previous experiences in a social context and may favor the persistence of left-handers in humans.

Cerebral lateralization of praxis in right‐ and left‐handedness: Same pattern, different strength

We aimed to investigate the effect of hand effector and handedness on the cerebral lateralization of pantomiming learned movements. Fourteen right-handed and 14 left-handed volunteers performed unimanual and bimanual tool-use pantomimes with their dominant or nondominant hand during fMRI. A left hemispheric lateralization was observed in the right- and left-handed group regardless of which hand(s) performed the task. Asymmetry was most marked in the dorsolateral prefrontal cortex (DLPFC), premotor cortex (PMC), and superior and inferior parietal lobules (SPL and IPL). Unimanual pantomimes did not reveal any significant differences in asymmetric cerebral activation patterns between left- and right-handers. Bimanual pantomimes showed increased left premotor and posterior parietal activation in left- and right-handers. Lateralization indices (LI) of the 10% most active voxels in DLPFC, PMC, SPL, and IPL were calculated for each individual in a contrast that compared all tool versus all control conditions. Left-handers showed a significantly reduced overall LI compared with right-handers. This was mainly due to diminished asymmetry in the IPL and SPL. We conclude that the recollection and pantomiming of learned gestures recruits a similar left lateralized activation pattern in right and left-handed individuals. Handedness only influences the strength (not the side) of the lateralization, with left-handers showing a reduced degree of asymmetry that is most readily observed over the posterior parietal region. Together with similar findings in language and visual processing, these results point to a lesser hemispheric specialization in left-handers that may be considered in the cost/benefit assessment to explain the disproportionate handedness polymorphism in humans.

Why are some people left-handed? An evolutionary perspective.

Since prehistoric times, left-handed individuals have been ubiquitous in human populations, exhibiting geographical frequency variations. Evolutionary explanations have been proposed for the persistence of the handedness polymorphism. Left-handedness could be favoured by negative frequency-dependent selection. Data have suggested that left-handedness, as the rare hand preference, could represent an important strategic advantage in fighting interactions. However, the fact that left-handedness occurs at a low frequency indicates that some evolutionary costs could be associated with left-handedness. Overall, the evolutionary dynamics of this polymorphism are not fully understood. Here, we review the abundant literature available regarding the possible mechanisms and consequences of left-handedness. We point out that hand preference is heritable, and report how hand preference is influenced by genetic, hormonal, developmental and cultural factors. We review the available information on potential fitness costs and benefits acting as selective forces on the proportion of left-handers. Thus, evolutionary perspectives on the persistence of this polymorphism in humans are gathered for the first time, highlighting the necessity for an assessment of fitness differences between right- and left-handers.

Socio‐economic status and handedness in two large cohorts of French adults

Left-and right-handers have coexisted since the Palaeolithic age. Hand preference is heritable. Moreover, there is extensive evidence of an association between left-handedness and several fitness costs. In this context, the persistence of the polymorphism is interesting. Here, we explore the associations between socio-economic status and handedness, analysing data from two large cohorts of adult men and women. Such associations are relevant to an evolutionary approach, as the socio-economic and the reproductive value are related. Our results partly support the hypothesis that left-handers have a socio-economic status advantage, countervailing the health issues. Although the models explain a small proportion of the variance observed, the frequency of left-handedness is significantly higher: (1) among women of higher educational level; (2) among categories of higher income; and (3) among individuals who have a higher position in the company. The importance of these findings for the evolution of the polymorphism of handedness is discussed.

Frequency-dependent performance and handedness in professional baseball players (Homo sapiens).

I used data on handedness and pitching and hitting performance in annual cohorts of professional baseball players (1957-2005) to test the hypothesis that handedness among pitchers was subject to negative frequency-dependent selection. As predicted by this hypothesis, right-handed pitchers were more successful (i.e., opposing batters hit more poorly against them) when they were relatively rare in the population. Contrary to the predictions of this hypothesis, however, left-handed pitchers were more successful when they were relatively common. Both right- and left-handed batters performed better in years dominated by right-handed pitchers, despite the fact that right-handed batters perform relatively poorly against right-handed pitchers. I suggest that batters form cognitive representations based on pitcher handedness, and that these representations are strengthened by repeated exposure or priming. When the pitcher handedness polymorphism is more balanced (e.g., 67% right-handed, 33% left-handed), these cognitive representations are less effective, which leads to decreased batting averages and improved performance by all pitchers. Furthermore, these cognitive representations are likely to be more critical to the success of right-handed hitters, who have reduced visuomotor skills relative to left-handed hitters.

Handedness and reproductive success in two large cohorts of French adults

Left- and right-handers in humans coexist at least since the Paleolithic, and this variation in hand preference has a heritable basis. Because there is extensive evidence of an association between left-handedness and several fitness costs, the persistence of the polymorphism requires an explanation. It is not known whether the frequency of left-handedness in Western societies is stable or not. If the polymorphism is at equilibrium and maintained by frequency dependence, it implies that the fitness of left-handers equals that of right-handers. On the contrary, if left- and right-handers have a different fitness, the polymorphism will evolve. Using two large cohorts of French adults (men and women), we investigated the relations between handedness and several estimators of the reproductive value: marital status, number of sexual partners (of the opposite sex), number of children, and number of grandchildren. Left-handers seem to have disadvantages for some life-history traits, such as marital status (for women) and number of children. For other traits, we observed sex-dependent interactions with socioeconomic status: for high-income categories, left-handed women report less sex partners and left-handed men have more grandchildren. These kinds of interactions are to be expected under the hypothesis that the polymorphism of handedness is stable.

Do left-handed children have advantages regarding school performance and leadership skills?

A handedness polymorphism exists in all human populations so far investigated, at least since the Palaeolithic. Handedness is a heritable trait, and a possible evolutionary explanation for the higher prevalence of right-handers is the association of left-handedness with some deleterious traits. In this context, the persistence of left-handers remains unexplained. Reproductive success has been shown to be dependent on cultural success in many human societies. This study is the first attempt to consider the importance of social status to evolutionary processes related to handedness. We conducted a survey in French public primary schools to look for a possible association between laterality, school performance, and leadership skills. This study revealed only weak associations, and a sex effect. For primary-school girls, we report a negative association between several indicators of left-sidedness and teacher's ratings of school performance and leadership skills. For primary-school boys, we report a positive association between left-handedness for writing and teacher's ratings of school performance. The present study did not reveal any strong relation between laterality and school performance, but has shown that general school performance is not independent of laterality, and that the effect is opposite in males and females. The main difficulty is now to define clearly the different intellectual functions involved.

Maintenance of handedness polymorphism in humans: a frequency-dependent selection model

Frequency-dependent selection is an important process in the maintenance of genetic variation in fitness. In humans, it has been proposed that the polymorphism of handedness is maintained by negative frequency-dependent selection, through a strategic advantage of left-handers in fighting interactions. Using simple mathematical models, we explore: (1) whether it is possible to predict the range of left-handedness frequencies observed in human populations by the frequency and the violence of fighting interactions; (2) the consequences of the sex differences in the probability of transmission of hand preference to offspring. We show that a wide range of values of the frequency of left-handers can be obtained with realistic changes of the parameters values. Our models reinforce the idea that negative frequency-dependence may have played a role in maintaining left-handedness in human populations, and provide further support for the importance of fighting interactions in the evolution of hand preference. Moreover, they suggest an explanation for the occurrence of left-handedness among women in this context, namely an indirect selective advantage through their male offspring.

Is there geographical variation in human handedness?

Right‐ and left‐handed individuals are present in all cultures. However, while it is known that handedness is a heritable trait, little is known about how handedness varies between populations—and without this knowledge, the significance of the left/right polymorphism is hard to interpret. We reviewed the literature to assess the extent of geographical variation of throwing or hammering handedness. These two tasks were chosen because they are present in all known cultures (unlike, for example, writing), and make sense within the context of several adaptive theories on the origin of laterality, or maintenance of handedness polymorphism, which state that tool or weapon manipulation are pivotal. A total of 81 samples were found with primary data on throwing or hammering handedness, spanning 14 countries and concerning more than 1,214,000 individuals studied between 1922 and 1998. A global logistic regression was performed to assess the significance of the country of the study, controlling for several potentially confounding variables (date of the study, sex and age of individuals). Country always had a significant effect, consistent with substantial geographical variation of throwing and hammering handedness. Curiously, left‐handedness frequency estimates for a given country were not always consistent across datasets, perhaps due to missing variables, such as educational level or socio‐economic status. Results are discussed in the context of the evolution of handedness and the significance of the current polymorphism.

Handedness: Neutral or adaptive?

Corballis seems to have not considered two points: (1) the importance of direct selection pressures for the evolution of handedness; and (2) the evolutionary significance of the polymorphism of handedness. We provide arguments for the need to explain handedness in terms of adaptation and natural selection.

Speech diagnosis of the states of norm and affective pathology

The coexistence of left- and right-handers is documented since prehistoric times, in all current ethnic groups, with some geographical frequency variations. Hand preference is influenced by genetic, hormonal, developmental, and cultural factors. Overall handedness is heritable, suggesting that this trait could evolve. Neutral models of evolution seem inadequate, as the generally low frequency of left-handers suggests some evolutionary costs. Data have suggested that left-handedness, as the rare hand preference, could represent an important strategic advantage in fighting interactions. The persistence of the polymorphism could thus be maintained by negative frequency-dependent selection. This fighting hypothesis, explaining the persistence of left-handers, applies only to situations where fighting abilities are directly linked to success (as in sports) or to Darwinian fitness, as in most past societies or in the few remaining groups not yet affected by Western colonization and a market economy. How the handedness polymorphism will evolve under the new modern conditions remains an interesting issue.

Local cortical blood flow, oxygen availability and electrical impedance responses to sensory stimulation

The coexistence of left- and right-handers is documented since prehistoric times, in all current ethnic groups, with some geographical frequency variations. Hand preference is influenced by genetic, hormonal, developmental, and cultural factors. Overall handedness is heritable, suggesting that this trait could evolve. Neutral models of evolution seem inadequate, as the generally low frequency of left-handers suggests some evolutionary costs. Data have suggested that left-handedness, as the rare hand preference, could represent an important strategic advantage in fighting interactions. The persistence of the polymorphism could thus be maintained by negative frequency-dependent selection. This fighting hypothesis, explaining the persistence of left-handers, applies only to situations where fighting abilities are directly linked to success (as in sports) or to Darwinian fitness, as in most past societies or in the few remaining groups not yet affected by Western colonization and a market economy. How the handedness polymorphism will evolve under the new modern conditions remains an interesting issue.