The use of bouts and frequencies in the evaluation of hand preferences for a coordinated bimanual task in chimpanzees (Pan troglodytes): An empirical study comparing two different indices of laterality.

Hand preferences for a coordinated bimanual task were assessed in 109 chimpanzees (Pan troglodytes). Hand preference was evaluated for 4 test sessions using bouts and frequencies of hand use to compare the sensitivity of each level of analysis in evaluating individual variation in handedness. Overall, significant population-level right-handedness was found using several different measures of hand use. Handedness indices based on bouts and frequencies were highly and significantly correlated. Moreover, hand preferences were consistent across tests despite efforts to situationally bias preference during each test. Taken together, these data do not support the view that bouts are a better level of analysis for evaluating hand preference. The results further suggest that hand preferences for coordinated bimanual actions are not influenced by situational factors and may reflect an inherent specialization of the left hemisphere for motor skill. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Note on hand use in the manipulation of joysticks by rhesus monkeys (Macaca mulatta) and chimpanzees (Pan troglodytes).

In a recent article MacNeilage, Studdert-Kennedy, and Lindblom (1987) proposed that nonhuman primate handedness may be contingent on the specific task requirements with visual–spatial tasks yielding left-hand preferences and fine motor tasks producing right-hand preferences. This study reports hand preferences in the manipulation of joysticks by 2 rhesus monkeys (Macaca mulatta) and 3 chimpanzees (Pan troglodytes). Reach data were collected on these same subjects and served as a basis for comparison with preference data for manipulation of the joystick. The data indicated that all 5 subjects demonstrated significant right-hand preferences in manipulating the joystick. In contrast, no significant hand preferences were found for the reach data. Reaction time (RT) data also indicated that the right could perform a perceptual–motor task better than the left hand in all 5 subjects. Overall, the data indicate that reach tasks may not be sensitive enough measures to produce reliable hand preferences, whereas tasks that assess fine motor control produce significant hand preferences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Visuospatial reaching preferences of common marmosets (Callithrix jacchus): An assessment of individual biases across a variety of tasks.

Using multiple measures of hand preference, the authors investigated lateralization at an individual level in 21 common marmosets. Despite showing group biases for sensory and communication functions, these same marmosets did not show a group bias in direction of lateralized hand use. Hand preferences were recorded on four novel reaching tasks requiring different levels of visual guidance and postural control. As found for simple food holding (with the same subjects), they displayed strong individual hand preferences but no group bias indicative of handedness. The strength of hand preference was influenced by task demands: stronger preferences were expressed when subjects adopted a suspended posture, and when "successful" versus "unsuccessful" foraging strategies were compared. Comparisons between visuospatial reaching and simple food holding preferences also revealed that half of the subjects displayed a division of function between the hands/hemispheres; subjects displayed opposing preferences in simple and visuospatial reaching, which would be beneficial for the performance of coordinated bimanual tasks. Given the apparent absence of a selective advantage for handedness, the authors suggest that hand preferences may reflect hemispheric dominance of other cognitive domains (i.e., temperament). (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Manual laterality in chimpanzees (Pan troglodytes) in complex tasks.

To determine the manual laterality of a sample of 24 chimpanzees, 4 problem apparatuses were used, the solution for which (obtaining a food item) required the use of 1 or both hands in sequential, simultaneous, or both sequential and simultaneous actions. The majority of the subjects showed significant and consistent hand preferences, especially in the actions that required a precision grip. The results obtained suggest the existence of factors linked to the specific characteristics of the task to be performed and to the ontogenetic maturation of the subject, which would influence the directionality and degree of the hand preference displayed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Comparative and familial analysis of handedness in great apes.

Historically, population-level handedness has been considered a hallmark of human evolution. Whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. This paper summarizes published data on handedness in great apes. Comparative analysis indicated that chimpanzees and bonobos show population-level right handedness, whereas gorillas and orangutans do not. All ape species showed evidence of population-level handedness when considering specific tasks. Familial analyses in chimpanzees indicated that offspring and maternal (but not paternal) handedness was significantly positively correlated, but this finding was contingent upon the classification criteria used to evaluate hand preference. Overall, the proportion of right handedness is lower in great apes compared with humans, and various methodological and theoretical explanations for this discrepancy are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Postural effects on manual reaching laterality in squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus).

Squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus) made unimanual food reaches from either a horizontal, quadrupedal posture or a vertical, clinging posture. No population-level handedness occurred in either species. However, in both species, directional lateral preferences weakly expressed for reaching from the stable quadrupedal posture were intensified in the vertical cling posture. This phenomenon, which is designated as soft handedness, may have been an evolutionary precursor to population-level handedness. Right or left turning by the squirrel monkeys before reaching closely predicted use of the right or left hand. However, the magnitude of the association decreased in the more highly lateralized vertical cling condition. This result suggests that as lateral hand preference increases, hand use may become increasingly independent of constraints from prior behavioral and environmental influences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preferences for simple reaching in juvenile chimpanzees (Pan troglodytes): Continuity in development.

Hand preferences were assessed in 51 chimpanzees (Pan troglodytes; 28 male and 23 female) ranging from 2 to 5 years of age. Simple reaching served as the measure of lateral bias in hand preference during 2 assessments separated by 1 year. A significant sex by hand preference interaction was found with a greater prevalence of right-handed males than females. No significant differences were found between age and either strength or direction of hand preference. A significant interaction was found between rearing and strength of hand preference. Mother-reared chimpanzees showed significantly greater strength in hand preference than nursery-reared chimpanzees. Finally, a significant positive correlation was found between tests of hand preference conducted over a 1-year interval. These data suggest that in chimpanzees hand preferences are established by 2 years of age and are stable throughout the juvenile developmental period. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Heritability of hand preference in chimpanzees (Pan troglodytes): Evidence from a partial interspecies cross-fostering study.

Heritability of hand preference was tested in a sample of 188 chimpanzees (Pan troglodytes). Hand preference was measured by coordinated bimanual actions, and concordance percentages were compared between parents and offspring and siblings. Among siblings, concordance percentages were compared for dyads in which both individuals were raised by chimpanzees, both were raised by humans, or 1 was raised in each environment. The results indicated population-level right hand preferences for coordinated bimanual actions. There were no significant associations in hand preference between parents and offspring. In full and maternal half siblings, concordance in hand preference was significantly greater than chance in mother and human-reared individuals but not in cross-fostered dyads. The cumulative results suggest that the direction of hand preference is heritable in chimpanzees but the mechanism of transmission is not genetic. Several environmental explanations are proposed to explain the findings, including the potential role of maternal cradling bias and in utero fetal position. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Grip morphology and hand use in chimpanzees (Pan troglodytes): Evidence of a left hemisphere specialization in motor skill.

Three experiments on grip morphology and hand use were conducted in a sample of chimpanzees. In Experiment 1, grip morphology when grasping food items was recorded, and it was found that subjects who adopted a precision grip were more right-handed than chimpanzees using other grips. In Experiment 2, the effect of food type on grasping was assessed. Smaller food items elicited significantly more precision grips for the right hand. In Experiment 3, error rates in grasping foods were compared between the left and right hands. Significantly more errors were made for the left compared with the right hand. The cumulative results indicate that chimpanzees show a left-hemisphere asymmetry in motor skill that is associated with the use of precision grips. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Lateral bias in infant chimpanzees (Pan troglodytes).

This study documents the presence, strength, and direction of lateralization in chimpanzees (Pan troglodytes) over the first 3 months of life. Nursery-reared chimpanzees (7 males and 5 females) were repeatedly assessed on a behavioral scale. Lateral bias was measured for 4 behaviors: hand-to-mouth, hand-to-hand, defensive grasp, and first step. Hand-to-mouth was significantly lateralized for the sample. Eight of the 10 chimpanzees that showed hand-to-mouth used the right hand. Lateral bias for defensive grasp was positively related to lateral bias both of first step and of hand-to-mouth. Lateral bias in hand-to-mouth was inversely related to lateral bias in hand-to-hand. Strength of lateralization increased as chimpanzees matured. These laterality effects in infant chimpanzees were expressed under conditions of emotional arousal. Moreover, degree of laterality may be a predictor of responsivity to stress. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Chimpanzees (Pan troglodytes) Are Predominantly Right-Handed: Replication in Three Populations of Apes.

Population-level right-handedness has historically been considered a hallmark of human evolution. Even though recent studies in chimpanzees (Pan troglodytes) have demonstrated population-level right-handedness for certain behaviors, some have questioned the validity and consistency of these findings by arguing that reported laterality effects are specific to certain colonies of apes and to those chimpanzees reared by humans. The authors report evidence of population-level right-handedness in 3 separate colonies of chimpanzees. Moreover, handedness in the 3 colonies was unrelated to the proportion of subjects that were raised by humans. This is the strongest evidence to date that population-level handedness is evident in chimpanzees and is not an artifact of human rearing. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Handedness for tool use correlates with cerebellar asymmetries in chimpanzees (Pan troglodytes).

Recent studies have shown that great ape species possess patterns of macrostructural neocortical asymmetries that are similar to those found in humans. However, little is known about the asymmetry of subcortical structures in great apes. To address this lack of data, the authors assessed left-right asymmetry of the anterior and posterior aspects of cerebellum from MRI brain scans of 53 chimpanzees (Pan troglodytes). No population-level bias was found for either the anterior or the posterior region of the cerebellum. However, a significant inverse association was found in the asymmetry quotients of the anterior and posterior regions, indicating that the cerebellum was torqued at the individual level. Additionally, handedness for tool use but not other measures was associated with variation in cerebellar asymmetries. Last, older chimpanzees had a smaller cerebellum after brain volume was adjusted for. The results are discussed in the context of brain changes in primate evolution related to tool use. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preferences for a coordinated bimanual task in 110 chimpanzees (Pan troglodytes): Cross-sectional analysis.

Hand preference for a coordinated bimanual task was assessed in a sample of 110 chimpanzees (Pan troglodytes). Subjects were presented with opaque tubes, the inside of which was coated with peanut butter. The hand and finger used to extract the peanut butter was recorded in 2 test sessions. A population right-hand bias was found. Juvenile and adolescent subjects were less lateralized than adults. All subjects primarily used their index finger to extract peanut butter. Use of the index finger was higher in adults and adolescents compared with juveniles. The results suggest that coordinated bimanual tasks (a) elicit strong hand preferences at the individual level and (b) elicit population level right-handedness. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Behavioral asymmetries of psychomotor performance in rhesus monkeys (Macaca mulatta): A dissociation between hand preference and skill.

Hand preferences were recorded for 35 rhesus monkeys as they manipulated a joystick in response to 2 computerized tasks. These preferences were then used to contrast 8 left- and 10 right-handed Ss on performance measures of hand skill. Individual hand preferences were found, but no significant population asymmetry was observed across the sample. However, the performance data reveal substantial benefits of right-handedness for joystick manipulation, as this group of monkeys mastered the 2 psychomotor tasks significantly faster than did their left-handed counterparts. The data support earlier reports of a right-hand advantage for joystick manipulation and also support the importance of distinguishing between hand preference and manual performance in research on functional asymmetries. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Examining the relationship between degree of handedness and degree of cerebral lateralization for processing facial emotion.

This paper examines the relationship between degree of handedness and degree of cerebral lateralization on a task of processing positive facial emotion in right-handed individuals. Three hundred and thirteen right-handed participants (157 women) were given two behavioral tests of lateralization: a handedness questionnaire and a chimeric faces test. Two further handedness measures were taken: familial lefthandedness and writing posture. Regression analysis showed that both degree of handedness and sex were predictive of degree of lateralization. Individuals who were strongly right-handed were also more strongly lateralized to the right hemisphere for the task. Men were more strongly lateralized than women. Data were reanalyzed for men and women separately. The relationship between handedness and lateralization remained for men only. Neither familial left-handedness nor writing posture were associated with cerebral lateralization for men or women. The results suggest a positive relationship between degree of handedness and degree of cerebral lateralization, and further that there is a sex difference in this relationship. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Right-handedness predominance in 162 baboons (Papio anubis) for gestural communication: Consistency across time and groups.

Whether nonhuman primates exhibit population level right manual bias remains a controversial topic. In captive baboons, population-level right-handedness has been reported for both coordinated bimanual actions and communicative gestures. However, some authors remain skeptical of these findings on both methodological and theoretical grounds. Here, we demonstrated the robustness and the consistency across time of the pattern of right-handedness for a species-specific communicative gesture in olive baboons (Papio anubis). First, we showed significant correlations in the 26 retested baboons for the measures of hand preferences between the first and the second session conducted 4 years later (2005–2009) by an observer blind to the previous handedness data. Second, the replication of the study in 96 novel individuals revealed a similar degree of population-level right-handedness than the one expressed in the first group of 66 subjects investigated in 2005. The implications of the findings are discussed within a theoretical framework about the origin of hemispheric specialization for language. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Assessment of shoulder dimensions and angles of porcelain bonded to metal crown preparations

This research examined hand preference for a bimanual task in 45 tufted capuchin (Cebus apella) and 55 rhesus macaque (Macaca mulatta) monkeys. Investigators presented subjects with plastic tubes lined with food and noted which hand the animals used to hold the tubes and which hand the animals used to remove the food. Several significant findings emerged from this investigation. First, rhesus macaques, but not tufted capuchins, exhibited a population-level bias toward use of the right hand (although the difference in direction of hand preference between species was not significant). Second, capuchins exhibited greater hand preference strength than did macaques. Third, among capuchins, but not among macaques, hand preference strength was greater for adults than for immatures. Finally, both species used their index digit to remove food most frequently when compared with other digits. Findings of hand preference direction and strength in this study were compared with other findings noted for chimpanzees which performed a bimanual tube task in a previous study. The authors conclude that using the same procedure to compare hand preference across species represents a powerful research tool that can lead to a more complete understanding of the evolution and ontogenesis of primate handedness.     

Sex and handedness effects on corpus callosum morphology in chimpanzees (Pan troglodytes).

Findings suggest that in humans, sex and hand preference may be associated with the size of the corpus callosum (CC). The authors measured CC morphology from MRIs in 67 chimpanzees (Pan troglodytes) to see whether similar effects were present in this species. Hand preference was assessed by performance on 4 tasks, and chimpanzees were classified as left-handed, right-handed, or ambidextrous. In a subsequent analysis, the chimpanzees were reclassified into 2 groups: right-handed and left-handed. The results revealed no sex difference in CC area, but significant effects of hand preference were found for several CC regions (rostrum body, anterior midbody, posterior midbody, isthmus, and splenium) and for overall CC size, with left-handed chimpanzees exhibiting significantly smaller CC measurements than right-handed chimpanzees. The results indicate that lateralized hand use in chimpanzees, as in humans, is associated with variation in CC size. (PsycINFO Database Record (c) 2010 APA, all rights reserved)