Hand preferences in captive orangutans (Pongo pygmaeus)

The strength of the evidence for population-level handedness in the great apes is a topic of considerable debate, yet there have been few studies of handedness in orangutans. We conducted a study of manual lateralization in a captive group of eight orangutans (Pongo pygmaeus) ranking the degrees of manual preference according to a defined framework. We analyzed five behavioral patterns: eat (one- and two-handed), make/modify tool, oral tool-use, and manual tool-use. Although some individuals showed significant manual preferences for one or more tasks, at the group-level both one-handed and two-handed eating, oral tool-use, and make/modify tool were ranked at level 1 (unlateralized). Manual tool-use was ranked at level 2, with four subjects demonstrating significant hand preferences, but no group-level bias to the right or left. Four subjects also showed hand specialization to the right or left across several tasks. These results are consistent with most previous studies of manual preference in orangutans. The emergence of manual lateralization in orangutans may relate to more complex manipulative tasks. We hypothesize that more challenging manual tasks elicit stronger hand preferences.     

Competitiveness,gender and handedness

We conduct an intercultural experiment in three locations on three different continents to elicit competitiveness and study whether individual differences in competitiveness are related to handedness. Being a “lefty” (i.e., having either a dominant left hand or a dominant left foot) is associated with neurological differences which are determined prenatally, and can therefore be seen as a proxy for innate differences. In large-scale data with incentivized choices from 3664 participants from India, Norway and Tanzania, we find a significant gender gap in competitiveness in all cultures. However, we find inconsistent results when comparing the competitiveness of lefties and righties. In north-east India we find that lefties of both genders are significantly more competitive than righties. In Norway we find that lefty men are more competitive than any other group, but women’s competitiveness is not related to handedness. In Tanzania, we find no relationship between handedness and the competitiveness of either gender. The merged data show weak evidence of a positive correlation between being a lefty and competitiveness for men, but no such evidence for women. Thus, our data provide suggestive but not robust evidence that individual and gender differences in competitiveness are partially determined by innate factors, where innate factors are proxied by the complex, prenatally shaped trait of handedness.     

Twisted growth and organization of cortical microtubules

In plants, directional cell expansion greatly contributes to the final shape of mature cells, and thus to organ architecture. A particularly interesting mode of cell expansion is helical growth in which the growth axis is continuously tilted either to the right or to the left as the cell grows. Fixed handedness of helical growth raises fundamental questions on the possible origin of left–right asymmetry. Twisting mutants of Arabidopsis thaliana offer unique opportunities to study the cellular basis of helical growth. Most of the twisting mutants with fixed handedness have been shown to have defects in microtubule functions, whereas mutants that twist in non-fixed directions appear to be defective in auxin response or transport. Good correlations have been found between the tilted growth direction and alignment of cortical microtubule arrays in twisting mutants with compromised microtubule functions. The present challenge is to understand how particular array patterns are organized during progression of the interphase in rapidly expanding cells. Molecular and cell biological studies on twisting mutants will lead to better understanding on how wild-type plant cells utilize the microtubule cytoskeleton to initiate and rigorously maintain straight growth. An erratum to this article can be found at     

Ant fishing by wild chimpanzees is not lateralised

Right-dominant handedness is unique and universal in Homo sapiens, suggesting that it is a highly derived trait. Our nearest living relations, chimpanzees, show lateralised hand preference when using tools, but not when otherwise manipulating objects. We report the first contrary data, that is, non-lateralised tool-use, for ant fishing as done in the Mahale Mountains of Tanzania. Unlike nut cracking, termite fishing, and fruit pounding, as seen elsewhere, in which most individuals are either significantly or wholly left- or right-biassed, ant fishers are mostly ambilateral. The clue to this exception lies in arboreality; all other patterns of chimpanzee elementary technology are done on the ground. Arboreal tool use usually requires not only that one hand be used to hold the tool, but also that the other hand gives postural support. When the supporting hand is fatigued, then it must be relieved by the other. Terrestrial tool use entails no such trading off. To test the hypothesis, we compared frequency of hand changing with the incidence of major hand support, and found them to be significantly positively correlated. The evolutionary transition from arboreality to terrestriality may have been a key enabler for the origins of human laterality.     

Handedness in wild chimpanzees

The debate over nonhuman primate precursors to human handedness is unsettled mainly due to lack of data, particularly on apes. Handedness in wild chimpanzees at the Taï National Park Côte d'Ivoire, has been monitored in four tasks. For the simple unimanual ones, reaching and grooming, adults use both hands equally (ambidextrous), while for the more complex unimanual wadge-dipping and the complex bimanual nut-cracking, adults are highly lateralized. These results support the hypothesis that lateralization increases with the complexity of the task. The lateralization is constant for years for each task but may vary in an individual with respect to different tasks. For nutcracking females are more lateralized than males. The ontogeny of handedness for nut-cracking shows many variations in the tendency to use one hand and in the side preferred, until at about 10 years of age, the individual achieves her adult handedness. No population bias toward one side exists in Taï chimpanzees. No heritability of handedness between mother and offspring was observed. Human and chimpanzees handedness are compared.     

Developmental origins of variation in human hand preference

Though right-handedness is a prominant characteristic within all human societies, a substantial and stable proportion of individuals are left-handed. Any comprehensive approach to the origin of variation in handedness must account for substantial evidence that left-handedness is associated with reduced fitness, neurodevelopmental disorders, and reduced neuroanatomical asymmetry. In this paper we investigate the hypothesis that developmental instability in early fetal development underlies variation in handedness. In two studies we note an increased incidence of minor physical anomalies (MPAs) and fluctuating asymmetries in both left-handers and extreme right-handers. Moreover, extreme right-handers were more apt to have left-handed parents than moderate right-handers. These data suggest that deviation from moderate right-handedness reflect imprecise expression of a near-universal design due to developmental instability. Preliminary attempts to elucidate the mechanisms underlying developmental instability suggest that both polygenic homozygosity and particular HLA alleles may be important factors. These observations are discussed with respect to current genetic theories of handedness and human evolution.     

Are gorillas right-handed or not?

Comparative data on laterality of function in primates are useful for elucidating its evolution, including its link to asymmetry of cerebral structure. Recently the gorilla (Gorilla gorilla, Pongidae) has been nominated as the other hominoid species closest toHomo sapiens in terms of handedness. This study aims to scrutinise the 22 accounts of gorillas' hand preferences. Numerous shortcomings exist in the published literature on the topic, so that a firm conclusion is not yet possible. It is not yet clear whether gorillas are right-preferent, left-preferent, or ambi-preferent, and additional data are needed.     

Genetic basis in motor skill and hand preference for tool use in chimpanzees (Pan troglodytes)

Chimpanzees are well known for their tool using abilities. Numerous studies have documented variability in tool use among chimpanzees and the role that social learning and other factors play in their development. There are also findings on hand use in both captive and wild chimpanzees; however, less understood are the potential roles of genetic and non-genetic mechanisms in determining individual differences in tool use skill and laterality. Here, we examined heritability in tool use skill and handedness for a probing task in a sample of 243 captive chimpanzees. Quantitative genetic analysis, based on the extant pedigrees, showed that overall both tool use skill and handedness were significantly heritable. Significant heritability in motor skill was evident in two genetically distinct populations of apes, and between two cohorts that received different early social rearing experiences. We further found that motor skill decreased with age and that males were more commonly left-handed than females. Collectively, these data suggest that though non-genetic factors do influence tool use performance and handedness in chimpanzees, genetic factors also play a significant role, as has been reported in humans.     

Factor Analysis of Multiple Measures of Hand Use in Captive Chimpanzees: An Alternative Approach to the Assessment of Handedness in Nonhuman Primates

We tested whether chimpanzee handedness could be characterized as either unidimensional or multidimensional when considered across multiple measures of hand use. We determined for each of 6 different tasks in a sample of 105 captive chimpanzees hand preferences, and subjected the individual hand preference scores to a factor analysis. Five of the 6 tasks loaded on two separate factors that accounted for 54% of the variance. To assess population-level handedness, we calculated handedness indices for the loadings on each factor, for the item loadings across all factors, and for all tasks including ones that did not load on any factor. There is significant population-level right handedness for all 4 indices, which suggests that chimpanzee handedness is multidimensional and not task specific.     

内蒙古汉、回、蒙古族扣手、交叉臂及惯用手的分析

利用对数线性模型分析了郑连斌、陆舜华等1996年在内蒙古调查的汉、回、蒙古族的3项人类遗传学指标（扣手、交叉臂、惯用手）,得出与其既有相同又有不同的结论。结果显示：（1）扣手与民族、性别都无关;（2）交叉臂与民族不相关,与性别相关;（3）惯用手、民族、性别三者存在三因子效应;（4）惯用手与扣手、惯用手与交叉臂分别存在相关性,而交叉臂与扣手之间相关性不显著。     

Hand preference in the Sifaka (propithecus verreauxi coquereli) during feeding in captivity

The Coquerel's sifakas were chosen for this study on hand preference because little is known about handedness in Indriidae. Fifteen Coquerel's sifakas were observed at the Duke University Primate Center as they fed on chopped fruit, vegetables, and primate chow. Analysis of age, sex, and hand preference indicated that the adult males both individually and as a group tended toward left-handedness. Adult females as a group did not show a trend in the direction of handedness. However, individual adult females showed consistent right- or left-hand preference. Younger sifakas tended toward ambipreference, suggesting that lateralization of hand preference is gradual, becoming more stable in adulthood. These findings suggest that sex and age may be strong indicators for lateralization of hand preference in Coquerel's sifakas. Duke University Primate Research Center Publication 292     

Brief communication: Effect of size biases in the coefficient of variation on assessing intraspecific variability in the prosimian skeleton

Lack of independence of data points or the pooling fallacy has been suggested as a potential problem in the study of handedness in nonhuman primates, particularly as it relates to whether hand use responses should be recorded as individual events or bouts of activity. Here, I argue that there is no evidence that the concept of statistical independence of data points or the pooling fallacy is a problem in the evaluation of population‐level handedness in previous studies in nonhuman primates. I further argue these statistical concepts have been misapplied to the characterization of individual hand preferences. Finally, I argue that recording hand use responses as bouts rather than events has no significant effect on reports of hand use in nonhuman primates and, in fact, may unintentionally bias hand use toward the null hypothesis. Several suggestions for improvement in the measurement and statistical determination of individual handedness are offered in the article. Am J Phys Anthropol 151:151–157, 2013. © 2013 Wiley Periodicals, Inc.     

Handedness in captive bonobos (Pan paniscus)

Species level right-handedness is often considered to be unique to humans. Handedness is held to be interrelated to our language ability and has been used as a means of tracing the evolution of language. Here we examine handedness in 3 captive groups of bonobos (Pan paniscus) comprising 22 individuals. We found no evidence for species level handedness. Conclusions that can be drawn from these findings are: (1) species level handedness evolved after the divergence of the Pan and Homo lineages; (2) inconsistent preferences may represent precursors to human handedness, and (3) Pan may have language abilities but these cannot be measured using handedness.     

Right‐handed fossil humans

Fossil hominids often processed material held between their upper and lower teeth. Pulling with one hand and cutting with the other, they occasionally left impact cut marks on the lip (labial) surface of their incisors and canines. From these actions, it possible to determine the dominant hand used. The frequency of these oblique striations in an array of fossil hominins documents the typically modern pattern of 9 right‐ to 1 left‐hander. This ratio among living Homo sapiens differs from that among chimpanzees and bonobos and more distant primate relatives. Together, all studies of living people affirm that dominant right‐handedness is a uniquely modern human trait. The same pattern extends deep into our past. Thus far, the majority of inferred right‐handed fossils come from Europe, but a single maxilla from a Homo habilis, OH‐65, shows a predominance of right oblique scratches, thus extending right‐handedness into the early Pleistocene of Africa. Other studies show right‐handedness in more recent African, Chinese, and Levantine fossils, but the sample compiled for non‐European fossil specimens remains small. Fossil specimens from Sima del los Huesos and a variety of European Neandertal sites are predominately right‐handed. We argue the 9:1 handedness ratio in Neandertals and the earlier inhabitants of Europe constitutes evidence for a modern pattern of handedness well before the appearance of modern Homo sapiens.     

Patterns of lateralized hand use in an arboreal primate, Simias concolor

Studies of hand use in nonhuman primates suggest that several species exhibit hand preferences for a variety of tasks. The majority of studies, however, focus on the lateralized hand use of captive nonhuman primate populations. Although captive settings offer a more controlled environment for assessing hand preferences, studies of wild populations provide important insights into how handedness is affected by natural environmental conditions and thus potential insights into the evolution of handedness. To investigate handedness in a population of wild nonhuman primates, we studied patterns of lateralized hand use during feeding in four simakobu monkeys (Simias concolor), an arboreal species inhabiting the Mentawai Islands, Indonesia. Our data show that individual variation in hand preferences for feeding existed among our study animals. In addition, each simakobu expressed a significant hand preference for supporting itself on a branch during feeding, an uncoordinated bimanual task. This bias was most prevalent when the branch used for support was a main branch rather than a terminal branch. When both hands were employed in a coordinated bimanual feeding activity (bimanual manipulation), only two subjects showed a significant bias for feeding. Our data suggest that these individuals are more likely to express significant hand preferences when feeding from stable, rather than precarious, positions within the canopy.     

Brief communication: Captive gorillas are right‐handed for bimanual feeding

Predominance of right‐handedness has historically been considered as a hallmark of human evolution. Whether nonhuman primates exhibit population‐level manual bias remains a controversial topic. Here, we investigated the hypothesis that bimanual coordinated activities may be a key‐behavior in our ancestors for the emergence and evolution of human population‐level right‐handedness. To this end, we collected data on hand preferences in 35 captive gorillas (Gorilla gorilla) during simple unimanual reaching and for bimanual coordinated feeding. Unimanual reaching consisted of grasping food on the ground, while bimanual feeding consisted of using one hand for holding a food and processing the food item by the opposite hand. No population‐level manual bias was found for unimanual actions but, in contrast, gorillas exhibited a significant population‐level right‐handedness for the bimanual actions. Moreover, the degree of right‐handedness for bimanual feeding exceeds any other known reports of hand use in primates, suggesting that lateralization for bimanual feeding is robust in captive gorillas. The collective evidence is discussed in the context of potential continuity of handedness between human and nonhuman primates. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Tryptophane-205 of human topoisomerase I is essential for camptothecin inhibition of negative but not positive supercoil removal

Positive supercoils are introduced in cellular DNA in front of and negative supercoils behind tracking polymerases. Since DNA purified from cells is normally under-wound, most studies addressing the relaxation activity of topoisomerase I have utilized negatively supercoiled plasmids. The present report compares the relaxation activity of human topoisomerase I variants on plasmids containing equal numbers of superhelical twists with opposite handedness. We demonstrate that the wild-type enzyme and mutants lacking amino acids 1–206 or 191–206, or having tryptophane-205 replaced with a glycine relax positive supercoils faster than negative supercoils under both processive and distributive conditions. In contrast to wild-type topoisomerase I, which exhibited camptothecin sensitivity during relaxation of both negative and positive supercoils, the investigated N-terminally mutated variants were sensitive to camptothecin only during removal of positive supercoils. These data suggest different mechanisms of action during removal of supercoils of opposite handedness and are consistent with a recently published simulation study [Sari and Andricioaei (2005) Nucleic Acids Res., 33, 6621–6634] suggesting flexibility in distinct parts of the enzyme during clockwise or counterclockwise strand rotation.     

Baseline cortisol levels and social behavior differ as a function of handedness in marmosets (Callithrix jacchus)

Population hand preferences are rare in nonhuman primates, but individual hand preferences are consistent over a lifetime and considered to reflect an individual's preference to use a particular hemisphere when engaged in a specific task. Previous findings in marmosets have indicated that left‐handed individuals tend to be more fearful than their right‐handed counterparts. Based on these findings, we tested the hypotheses that left‐handed marmosets are (a) more reactive to a social stressor and (b) are slower than right‐handed marmosets in acquiring a reversal learning task. We examined the hand preference of 27 male and female marmosets (ages of 4–7 years old) previously tested in a social separation task and a reversal learning task. Hand preference was determined via a simple reaching task. In the social separation task, monkeys were separated from their partner and the colony for a single 7‐hr session. Urinary cortisol levels and behavior were assessed at baseline, during the separation and 24 hr postseparation. Hand preferences were equally distributed between left (n = 10), right‐handed (n = 10), and ambidextrous (n = 7) individuals. The separation phase was associated with an increase in cortisol levels and behavioral changes that were similar across handedness groups. However, cortisol levels at baseline were positively correlated with right‐handedness, and this relationship was stronger in females than in males. In addition, the occurrence of social behaviors (pre‐ and postseparation) was positively correlated with right‐handedness in both sexes. Baseline cortisol levels did not correlate significantly with social behavior. Acquisition of the reversals was poorer in females than males but did not differ as a function of handedness. We conclude that (a) both stress reactivity and cognitive flexibility are similar across handedness groups and (b) left‐handers exhibit less social behavior and have lower basal cortisol levels than ambidextrous and right‐handed subjects. The underlying causes for these differences remain to be established.     

On the handedness of Japanese monkeys

Forty-one monkeys living on the Koshima Island were tested for handedness by throwing a peanut 10 or 20 times toward each subject and recording which hand was used to pick it up. Handedness was judged at the 5% level of confidence. Animals showing no preference for the left or right hand after 20 trials were considered to be ambidextrous. The following distribution of handedness was found: left-handed, 17 (41%); right-handed, 8 (20%); ambidextrous, 16 (39%). These findings were compared with other studies on the handedness of Japanese monkeys. Handed monkeys outnumber the ambidextrous ones and the left-handed monkeys outnumber the right-handed ones. The characteristic distribution of handedness was shown to be irrespective of troop. By looking over the data from these different sources, the proportions of the left-handed, the ambidextrous, and the right-handed in Japanese monkeys were estimated at 39%, 33%, and 28% respectively. The method used in this study was compared withKawai's catching behavior and reasons for disagreement between judgments of handedness were discussed.