Origins and functions of laterality: interactions of motoric systems

To evaluate lateral motoric bias in response systems at different levels of the neuraxis and assess the extent of interaction between these levels in the smalleared bushbaby ( Otolemur garnettii ), 27 animals were tested for lateral bias in hand use and whole-body turn bias in two postural conditions. Subjects retrieved mealworms quadrupedally by reaching downwards into glass jars and bipedally by reaching upwards to baited straws. Eye bias was assessed separately. Behaviours were scored from videotape. Two subgroups were identified: SHIFTERS changed hand preference with posture and had correlations of hand/eye bias in quadrupedal posture and of hand/turn bias, with more bimanual reaching, in bipedal posture; NONSHIFTERS were consistent in hand preference and more strongly lateralised in reach and turn than SHIFTERS. Subgroups did not differ in reach efficiency. Results are interpreted to support the value of the analysis of motoric levels and their interactions in the study of the evolution of laterality. Assuming natural selection for coordinated and targeted behaviours to be the source of lateralisation, several proposals in support of a motoric theory of laterality origins and functions are advanced.     

Age-related decline in lateralised prey capture success in Garnett's bushbaby (Otolemur garnettii)

We examined differences in prey capture success when reaching for moving prey with the preferred and non-preferred hand (as determined previously using stationary food items) in 12 Garnett's bushbabies (Otolemur garnettii). Hand preference was determined by a test of simple reaching for stationary food items. We assessed both the frequency of hand use and success rates for each hand in capturing live mealworms. We also examined the effect of age on overall prey capture success. Subjects were individually presented with live mealworms in a cup partially filled with a cornmeal medium. The preferred hand was used significantly more often than the non-preferred hand to obtain the moving prey; however, no differences were found in the frequency of usage of the left vs the right hand. Furthermore, there were no differences in the success rates of the left vs the right hand, nor the preferred vs the non-preferred hand. There was a significant negative correlation between age and prey capture success. These data suggest that age, rather than preferred hand, may be the most relevant factor in the bushbabies’ prey capture success.     

Posture and reaching in tufted capuchins (Cebus apella)

This research investigated the effects of posture on lateral bias for food reaching in tufted capuchin monkeys ( Cebus apella ) by comparing hand preferences for quadrupedal and bipedal reaching. Several findings of this investigation warrant discussion. First, we found a population-level bias towards use of the right hand for bipedal reaching but not for quadrupedal reaching. Second, adults exhibited a greater right-hand preference for bipedal reaching than did immatures. Third, subjects showed a greater right-hand preference, and a greater strength of preference independent of direction, for bipedal reaching than for quadrupedal reaching. Fourth, we found a significant positive relation between the direction of hand preference for quadrupedal and bipedal reaching. We believe that capuchins provide an alternative primate model to chimpanzees for the evolution of human bipedalism and right-handedness. One implication of this model is that righthandedness emerged in hominids prior to extensive expansion of brain size and elaboration of material culture.     

The effect of situational factors on hand preferences for feeding in 177 captive chimpanzees (Pan troglodytes)

This study examined the effect of situational factors on hand use for feeding in a sample of captive chimpanzees. Lateral bias in hand use was tested in biased and unbiased testing circumstances to assess strength and consistency in hand preference. For both unbiased and biased testing conditions, population-level right hand preferences were found for the sample. In the biased condition, subjects were more likely to overcome positional factors in order to feed with their right hand contrasted with the left. Overall, hand use in the biased and unbiased testing conditions was significantly positively correlated. In terms of strength of hand use, juveniles were found to be less lateralized than sub-adults and adults. Moreover, juvenile females were found to be more lateralized than juvenile males. Taken together, the data suggest that chimpanzee hand preferences for feeding are not constrained by situational factors and are relatively consistent in biased and unbiased testing conditions.     

Turning bias and lateral dominance in a sample of able-bodied and amputee participants

Turning bias is the tendency to turn towards a given direction. Conflicting results from previous studies suggest that a number of factors may influence turning direction. The aim of this study was to determine if biomechanical asymmetries influence turning bias. A total of 100 able-bodied participants, and 30 trans-tibial amputees who, by definition, possess a functional asymmetry, volunteered to participate in the study. The right hand and right foot were significantly dominant for the able-bodied sample. Able-bodied participants showed a significant turning preference towards the left, which was opposite to the dominant hand and foot. The amputees were significantly right-hand dominant and the side of the amputation influenced foot dominance. The amputee sample showed no preferred turning direction. Turning bias indices in the amputee sample were not significantly associated with handedness, footedness, side of amputation, or dominance prior to amputation. The lack of a preferred direction of turn in the amputee sample suggests that biomechanical asymmetries can influence turning bias.     

Turning bias and lateral dominance in a sample of able-bodied and amputee participants

Turning bias is the tendency to turn towards a given direction. Conflicting results from previous studies suggest that a number of factors may influence turning direction. The aim of this study was to determine if biomechanical asymmetries influence turning bias. A total of 100 able-bodied participants, and 30 trans-tibial amputees who, by definition, possess a functional asymmetry, volunteered to participate in the study. The right hand and right foot were significantly dominant for the able-bodied sample. Able-bodied participants showed a significant turning preference towards the left, which was opposite to the dominant hand and foot. The amputees were significantly right-hand dominant and the side of the amputation influenced foot dominance. The amputee sample showed no preferred turning direction. Turning bias indices in the amputee sample were not significantly associated with handedness, footedness, side of amputation, or dominance prior to amputation. The lack of a preferred direction of turn in the amputee sample suggests that biomechanical asymmetries can influence turning bias.     

A group test for the assessment of performance between the hands

A group performance test of handedness was administered to 1556 undergraduates. This test required subjects to place dots in circles as rapidly as possible. The test was found to be reliable, and to correlate with hand preference. Both hand performance and hand preference measures are skewed in the population as a whole, but the distributions can be adequately fitted by two normal curves, one with a right bias and one with a left bias. These findings suggest that left-handers are a distinct subgroup of the population.     

Left-handedness and longevity in primates.

In the present research we used the rhesus macaque to model the decline in the incidence of left-handedness that has been noted in the aged human population. We found a significant group-level bias towards use of the left hand among young-adult macaques, and a significant group-level bias towards use of the right hand among aged macaques. The distribution of hand preference across age classes cannot be explained through simple elimination of left-handed subjects in the aged population. Rather, our data are consistent with a maturational view positing increased use of the right hand with increased age. Similar findings across phylogenetically diverse primate taxa suggest that this phenomenon is an evolutionarily ancient trait. Greater use of the left hand in male versus female macaques indicates that sex differences in hand preference are also deeply rooted in our primate origins.     

Lateralized preferences and right-left discrimination in the monkey: choice of mirror-image shape, position of recovered food and hand usage.

Rhesus monkeys were tested for lateral preferences during mirror-image shape discrimination and food recovery from a row of pellets. Position of recovered food was unrelated to hand preference or mirror-image preference. There was no significant difference in the number of individuals preferring right and left lateral mirror-image shapes. Individuals tended to retain the same mirror-image preference to the same degree upon repeated testing. Animals of greater weight had stronger preferences. In all these respects, the features of mirror-image preference resembled those previously described for hand preference during a simple reaching task. The strength of different lateral preferences expressed by each animal were not significantly related thus failing to support the concept that asymmetric bodily responses can serve as cues for right-left discrimination in the environment. However, when the direction as well as strength of preference was considered, there appeared to be a tendency for animals to prefer the mirror-image shape whose reward bias was opposite to that of the preferred hand. This finding might have been due to a positional bias though there was evidence against this possibility.     

Handedness for eating in gorillas

Observations of spontaneous reaching for food in 31 captive lowland gorillas found a full range of hand preferences between strong left and strong right, with most animals showing intermediate levels of preference. There was a high degree of consistency between observations made on different occasions for the same animal, showing that degrees of relative preference tend to be stable in individuals. The findings agree with the majority of previous studies of apes and other primates in showing no species bias to one side, with about half the animals left and half right preferent.     

Hand preferences for bimanual coordination in 29 bonobos (Pan paniscus)

Brain lateralization has long been thought to be unique to humans. To investigate the origins and functions of this feature, researchers study behavioural laterality in other animals. Despite a substantial database, manual laterality in non-human primates remains a controversial topic. We give here a review of the main findings on manual preference in great apes. This article presents data on hand preferences for a bimanual coordination in 29 bonobos (Pan paniscus). The study aims to provide data on manual laterality for a complex bimanual task in this very interesting and rarely studied species. Hand preferences were assessed using the 'tube task'. This task has been used with other species, which allows reliable data comparisons. The task requires a bimanual coordinated precise action: the subject holds the tube with one hand while reaching for food inside with the other hand. As a complex task, this measure has been shown to be efficient in revealing hand preferences. It has revealed group-level right bias in chimpanzees. Bonobos had never been tested. We recorded both independent bouts (counting only the first pattern of a sequence of identical actions) and frequency (counting every action). The bonobos exhibited strong hand preferences. With frequency, 11 bonobos were classified as right-handed, 15 were left-handed and 3 had no preference. With bouts, 8 bonobos were right-handed, 9 were left-handed and 12 had no preference. No group-level bias appeared. The results are discussed in relation with previous findings and theories on brain lateralization.     

Manual specialisation and tool use in captive chimpanzees (Pan troglodytes): the effect of unimanual and bimanual strategies on hand preference

Hand preference for tool use was assessed in a sample of captive chimpanzees ( Pan troglodytes ). Whether the subjects solved the tool task using either a unimanual or coordinated bimanual strategy was manipulated in the chimpanzees. No population-level hand preference was found for tool use when unimanual strategies were used by the chimpanzees. However, a population-level right-hand bias was found when coordinated bimanual actions were required of the chimpanzees. A significant correlation was found in hand use for the two hand preference testing conditions. Neither sex nor rearing was found to significantly affect the direction or strength in hand preference. These results may explain discrepancies in hand preference reported in captive and wild chimpanzees with regard to tool use and other manual activities.     

Dorsal visual cortex activity elicited by posture change in a visuo-tactile matching task

To investigate the process of crossmodal spatial recognition, we examined the effect of posture change on the recognition of a tactile stimulus position. The task was to judge whether a visual and a tactile stimulus, presented to the left or right, were on the same or different sides while subjects crossed or uncrossed their hands. Under a condition which removed the effect of response bias to the left and right, the dorsal visual cortex (area 18/19) and the precuneus were more activated in the crossed hands condition. The dorsal visual cortex activation suggests that the activity of brain areas classically considered to be visual cortex is affected by posture change, and reflects the reciprocal process across different modalities in spatial recognition.     

The influence of classical dance training on preferred supporting leg and whole body turning bias

A rightward turning bias has been more frequently noted during adult classical dance practice than during spontaneous rotations. Training could play a role in inducing a preferred direction. We observed the preferred direction for executing four spontaneous whole-body full turns (pirouettes), with eyes open or closed, in pre-pubertal untrained girls and classical dance students. Of untrained girls, 58% showed a leftward turning bias (LTB) and 42% a rightward turning bias (RTB), independently of vision, lateral preferences, and supporting leg. Only one dancer showed a consistent LTB while the majority showed a RTB, with a tendency to use the left leg to turn towards the right. These results suggest that the role of the vestibular and visual systems is minimal for untrained girls, and suggest a training influence for dancers. The dance students' choice of a supporting leg for turning may exploit some biomechanical properties facilitating the pirouette.     

The influence of classical dance training on preferred supporting leg and whole body turning bias

A rightward turning bias has been more frequently noted during adult classical dance practice than during spontaneous rotations. Training could play a role in inducing a preferred direction. We observed the preferred direction for executing four spontaneous whole-body full turns (pirouettes), with eyes open or closed, in pre-pubertal untrained girls and classical dance students. Of untrained girls, 58% showed a leftward turning bias (LTB) and 42% a rightward turning bias (RTB), independently of vision, lateral preferences, and supporting leg. Only one dancer showed a consistent LTB while the majority showed a RTB, with a tendency to use the left leg to turn towards the right. These results suggest that the role of the vestibular and visual systems is minimal for untrained girls, and suggest a training influence for dancers. The dance students’ choice of a supporting leg for turning may exploit some biomechanical properties facilitating the pirouette.     

Focal transcranial magnetic stimulation and response bias in a forced-choice task.

The effects of transcranial magnetic stimulation were studied on the performance of a warned, forced-choice response time task by normal adults. The task consisted of extension of the index finger in response to the click produced by the discharge of the magnetic coil (go-signal). The subjects were asked to choose the right or left finger only after the go-signal was delivered. Single magnetic stimuli were delivered to the prefrontal or motor area, and in the control situation, away from the head. Magnetic stimulation affected hand preference only when it was delivered to the motor area. With stimulation of this area, subjects more often chose the hand contralateral to the site stimulated with response times that were mainly less than 200 ms. With longer response times (between 200 and 1100 ms), magnetic stimulation had no effect on hand preference regardless of the site stimulated. Stimulation of prefrontal areas yielded results similar to the control situation. These results suggest that response bias in this paradigm is caused by an effect of magnetic stimulation on neural structures within, or closely related to, the motor areas of the brain. Although the response bias was clear and predictable, the subjects were unaware of its existence. It is possible to influence endogenous processes of movement preparation externally without disrupting the conscious perception of volition.     

Hand preference by black-and-white snub-nosed monkeys (Rhinopithecus bieti) in captivity: influence of tasks and sexes

Task complexity (Fagot & Vauclair 1991), bimanual complementary role differentiation (Uomini 2009), and the obligate use of a particular hemisphere (Rogers, 2009) have been proposed to explain why hand preferences in non-human primates are often influenced by tasks. We examined how tasks (reaching, carrying, extractive foraging, and object manipulation) and gender influenced hand preference in 11 adult black-and-white snub-nosed monkeys (Rhinopithecus bieti) out of a total of 13 known adult captive individuals of this species. A logistic regression was used to analyse 2556 bouts of binary left- vs right-hand use data. The explanatory variables are tasks, gender, and the interaction of tasks and gender. Hand preference is influenced by the task, in that subjects used the right hand significantly more often for extractive foraging and object manipulation than for reaching and carrying. We also found a significant interaction of sex and task: males used the left hand significantly more often than females for reaching and carrying, respectively, but not for extractive foraging or object manipulation. This is the first study on hand preference in R. bieti. As predicted, the hand preference in R. bieti is not a fixed property of the species or sexes but depends on the task.     

Hand preference by black-and-white snub-nosed monkeys (Rhinopithecus bieti) in captivity: Influence of tasks and sexes

Task complexity (Fagot & Vauclair 1991), bimanual complementary role differentiation (Uomini 2009), and the obligate use of a particular hemisphere (Rogers, 2009) have been proposed to explain why hand preferences in non-human primates are often influenced by tasks. We examined how tasks (reaching, carrying, extractive foraging, and object manipulation) and gender influenced hand preference in 11 adult black-and-white snub-nosed monkeys (Rhinopithecus bieti) out of a total of 13 known adult captive individuals of this species. A logistic regression was used to analyse 2556 bouts of binary left- vs right- hand use data. The explanatory variables are tasks, gender, and the interaction of tasks and gender. Hand preference is influenced by the task, in that subjects used the right hand significantly more often for extractive foraging and object manipulation than for reaching and carrying. We also found a significant interaction of sex and task: males used the left hand significantly more often than females for reaching and carrying, respectively, but not for extractive foraging or object manipulation. This is the first study on hand preference in R. bieti. As predicted, the hand preference in R. bieti is not a fixed property of the species or sexes but depends on the task.     

Childrens’ left-turning preference is not modulated by magical ideation

The literature on human turning preferences is inconsistent. While the few studies with children below 14 years of age uniformly describe an overall left-turning (counterclockwise) tendency, a recent Internet study with more than 1500 adults found a right-sided (clockwise) bias. We set out to investigate spontaneous turning behaviour in children age 5–3 years and, based on neuropsychiatric work in adults, also explored a potential association with magical thinking. Findings indicated a clear left-turning preference, independent of a participant's sex and handedness. Whether a child responded a question about the existence of extrasensory communication in the affirmative or not was unrelated to direction and size of turning bias and lateral preference. Our results are consistent with a left-sided turning preference reported for children, but in opposition to the clockwise bias recently described in a large-scale study with adults. Whether they point to a maturational gradient in the preferred direction of spontaneous whole-body rotation or rather to a lack of comparability between measures used in observational versus Internet-based studies remains to be further investigated. Regarding a purported association between body turns and magical thinking, our study is preliminary, as only one single question was used to probe the latter.     

Posture and laterality in the bushbaby (Galago senegalensis)

A colony of adult bushbabies was assessed for hand preference in order to determine what the proportion of preference for the left and right might be for the population and whether hand preference for individuals was reliable. 25 animals were tested in an apparatus demanding a vertical stance. The population bias was expressed in a distribution skewed toward a left-hand bias. With 16 animals available for retesting, the test-retest reliability coefficient was 0.651, significant at the 0.01 level. When 12 of the original subjects were tested in an apparatus demanding a quadrupedal stance, hand preference shifted to a bimodal distribution. Test-retest reliability was 0.864, significant at the 0.01 level. Significant correlations were not found between two types of test nor between the two retests. Neither sex nor length of laboratory residence served to predict hand preference. These results were discussed with the view that the postural adjustments required by a bipedal stance may have shaped the development of handedness in humans.