Hand Preference for a Novel Bimanual Coordinated Task During Termite Feeding in Wild Western Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Although the level of handedness in humans varies cross-culturally, humans are generally described as right-handed, which has been considered a uniquely human trait. Recently, captive chimpanzees (Pan troglodytes) have been shown to exhibit right-hand preference when performing bimanual but not unimanual tasks. Less clear is whether this pattern also occurs in wild chimpanzees and other African apes. Using videos (N = 49) of six wild western gorillas (Gorilla gorilla gorilla) feeding on termites at the Mondika Research Center (Republic of Congo), we tested whether they exhibit hand preference when performing unimanual, i.e., reaching for termite mound pieces; bimanual, i.e., “termite tapping”: rhythmically shaking a piece of termite mound with the dominant hand and collecting the termites in the other hand tasks; or hand transfer prior to bimanual tasks, i.e., transferring a piece of termite mound from one hand to the other. All individuals exhibited exclusive hand preference when performing the bimanual tasks, with five of six gorillas preferring the right hand. Conversely, most individuals did not show any manual preference during the unimanual task. In addition, hand preference during hand transfer revealed clear hand dominance of similar strength and direction of those shown for the bimanual task, suggesting that this measure is as sensitive as the bimanual task itself. Thus, we propose “termite feeding” as a novel task to be considered in future hand-preference studies in wild western gorillas. Our results are in concordance with those for chimpanzees and captive gorillas showing hemispheric specialization for bimanual actions in apes.     

Hand preferences for a haptic searching task by tufted capuchins (Cebus apella)

We tested the effects of a haptic search task on hand preferences in capuchins(Cebus apella) and compare this situation to a visual by guided reaching task. In the haptic task, 21 monkeys searched for sunflower seeds on the top or side surfaces of 12 objects. A left-hand preference emerged at the group level, suggesting a greater involvement of the right hemisphere. The percentage of preferred hand usage and the direction of the preference were influenced by both sex and age of the subjects: adult males tended to be less lateralized than the other groups of subjects were. Shape had an insignificant effect on the direction of hand preferences or on the percentage of preferred hand use. No lateral bias emerge in the visually guided reaching task, and the percentage of left-hand usage fell significantly across tasks, demonstrating that the haptic demands of the task enhance the use of the left hand at the group level. We discuss these results with regard to current theories on manual lateralization in nonhuman primates.     

Hand Preferences in Different Tasks by Tufted Capuchins (Cebus apella)

We examined hand preferences in 25 tufted capuchins (Cebus apella) in three tasks. The hole task involved a single action of reaching for food in a hole. The horizontal panel and the vertical panel tasks required the alignment of two apertures, by moving or lifting a panel, to reach for food in a hole. We found a significant group-level right-hand preference for reaching actions in the hole and in the horizontal panel tasks, but not in the vertical panel task, in which the food retrieval implied the complementary use of both hands. No significant hand bias emerged for moving or lifting actions with high visuospatial components. There is a stronger hand preference in more complex manual activity—coordinated bimanual hand use for food retrieval—than in other unimanual measures. We discuss the results in the context of previous reports on primate laterality.     

Population-Level Right Handedness for a Coordinated Bimanual Task in Chimpanzees: Replication and Extension in a Second Colony of Apes

The purpose of this study was to evaluate the reliability of previously published findings on hand preferences in chimpanzees by evaluating hand use in a second colony of captive chimpanzees. We assessed hand preferences for a coordinated bimanual task in 116 chimpanzees (Pan troglodytes) at the University of Texas M. D. Anderson Cancer Center and compared them to previously published findings in captive chimpanzees at the Yerkes National Primate Research Center. The new sample showed significant population-level right handedness, which is consistent with previously published findings in the Yerkes chimpanzees. Combined data on the 2 chimpanzee colonies, revealed a significant effect of rearing history on hand preference, with wild-caught chimpanzees showing less right-handedness than captive-born mother-reared chimpanzees. We discuss the results in terms of the role of early environment on the development of laterality.     

Hand preference in a captive island group of chimpanzees (Pan troglodytes)

Morphological cerebral asymmetries in chimpanzee brains, similar to those found in humans, in whom they are associated with speech and handedness, suggest the possibility of functional lateralization in the chimpanzee. This possibility was investigated by examining hand preferences in an island group of five chimpanzees on a series of unimanual and bimanual tasks that are diagnostic of human hand and cerebral dominance. Each subject was tested in a double compartment cage on three unimanual nonsequential, three unimanual sequential, and three bimanual coordination tasks. One of the three unimanual sequential tasks was a bar-press task that is analogous to the commonly used human finger-tapping task. For the unimanual tasks, exclusive of the bar-press, the chimpanzees showed a highly individualistic pattern of hand preference that did not change as a function of task complexity. On the bar-press task, four of five subjects produced higher rates with one hand compared to the other; however, relative hand performance on this task was unrelated to hand preference on the other unimanual tasks. For the group of subjects, performance rates did not differ between the left and right hands; however, a practice effect was observed for the right hand in all subjects. The bimanual tasks also revealed a complex pattern of individual handedness, with no trends apparent for the group as a whole. Consistent with previous findings, the results from these tests on this group of five chimpanzees suggest that cerebral morphological asymmetries in the chimpanzee are not associated with motor dominance as reflected in handedness.     

Manual Laterality for Simple Reaching and Bimanual Coordinated Task in Naturalistic Housed <Emphasis Type="Italic">Pan troglodytes</Emphasis>

Researchers have shown renewed interest in the study of manual lateralization in chimpanzees. Currently there is no consensus confirming the presence or absence of manual dominance at a species level, mainly for populations in the wild and in semicaptivity. We aimed to evaluate the manual laterality in a group of chimpanzees in an intermediate setting (semicaptivity) via 2 tasks: one simple and unimanual (simple reaching) and the other complex and bimanual (tube task). We replicated the same experiments from Hopkins in a new and different sample of chimpanzees. In simple reaching, the hand is used to gather food and the type of grip and the posture are evaluated. The tube task assesses the hand used to extract food from the tube and the method of extraction (digital or instrumental). Through the handedness index we observed that the subjects show clear and strong individual preferences for both tasks (100% lateralized subjects in the tube task; 86% in simple reaching), although we did not detect population preferences for any of the tasks. However, considering both tasks jointly (multiple evaluation), it was possible to detect, for the first time, skilled manual dominance at a group level in semicaptive chimpanzees in one multitask index and borderline significance in a second multitask index.     

Brief communication: Hand preference for bimanual and unimanual feeding in captive gorillas: extension in a second colony of apes

Right-hand dominance is widely considered to be a uniquely human trait. Whether nonhuman primates exhibit similar population-level hand preferences remains a topic of considerable debate. Despite extensive research focusing on laterality in nonhuman primates, our interpretation of these studies is limited due to methodological issues including the lack of a common measure of hand preference and the use of tasks that may not be reliable indicators of handedness. The use of consistent methods between studies is necessary to enable comparisons within and between species and allow for more general conclusions to be drawn from these results. The present study replicates methods used in recent research reporting population-level right-handedness in captive gorillas (Meguerditchian et al.,2010). Observational data were collected on hand preference for unimanual and bimanual feeding in 14 captive western lowland gorillas (Gorilla gorilla gorilla). Individual-level preferences were found, primarily for bimanual feeding; however, the data reveal no group-level directional bias (contra Meguerditchian et al.). Like the study by Meguerditchian et al. (2010), though, bimanual feeding revealed significantly stronger hand preferences than unimanual reaching, and age, sex, group membership, or rearing history had no effect on hand preference. Finally, variations in diet and corresponding grip type between studies suggest that hand preferences may vary across bimanual tasks depending on grip morphology. This study aims to contribute to our existing knowledge of primate laterality by increasing the number of individuals investigated using methods that allow for comparisons with similar research.     

Does the Complexity of the Task Influence Manual Laterality in De Brazza’s Monkeys (Cercopithecus neglectus)?

We assessed the manual preferences of 12 De Brazza's monkeys ( Cercopithecus neglectus ) in spontaneous feeding situations and in two different coordinated bimanual tasks that were not visually guided. We recorded the hand used by each subject for 22 spontaneous activities, hand and digits use while extracting peanut butter from a hollow tube (tube task) and the hand used to extract candies from hanging plastic balls (ball task). Spontaneous activities revealed individual manual preferences but no population-level biases. For both experimental tasks, all subjects were lateralized in their hand use. We found a left bias at the group level for the tube task, but no group-level asymmetry for the ball task. Experimental tasks induced greater strength of laterality than did spontaneous activities. Although the size of our sample did not allow us to draw any conclusions concerning manual preference at the population level, this study stresses the importance of coordinated bimanual tasks to reveal manual laterality in non-human primates.     

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.     

Sex difference in chimpanzee handedness

Chimpanzees at Mahale, Tanzania, show strong individual hand preferences when they use bimanual actions in processing the fruit of Saba florida and Citrus lemon. The direction of hand preference differs between the sexes: most males are left-handed, whereas most females are right-handed. Monkeys and apes are considered to lack "handedness," in the sense of a population mode of left- or right-hand preference; they are normally ambidextrous. Indeed, strong individual preferences were previously seldom found in natural tasks. We propose that lateralization of manual actions becomes advantageous in bimanual tasks, which involve role differentiation between the hands and a need to combine power and precision. If the pattern of lateralization found here reflects the ancestral state, common to chimpanzees and humans, this may explain why, in modern humans, women tend more strongly to be right-handed than men, who include a larger minority of left-handers.     

Hand preferences on unimanual and bimanual tasks in Tonkean macaques (Macaca tonkeana)

This is the first study to examine hand preferences in Tonkean macaques on a bimanual task. One of our objectives was to continue the move toward greater task standardization, in order to facilitate comparisons between species and studies on handedness. The main aim was to test and determine task robustness, by varying intra‐task complexity. To this end, we administered several different tasks to the subjects: two unimanual tasks (grasping task featuring items of different sizes) and three coordinated bimanual tasks (tube task involving different materials, weights, and diameters). Although we found no significant hand preference in either task at the group level, the macaques were more strongly lateralized for small items than for large ones in the unimanual grasping task. Moreover, the absence of a correlation between these two versions of the unimanual task confirmed the weakness of this grasping task for assessing handedness. Regarding the bimanual tube task, no difference was found between the three versions in either the direction or the strength of hand preference. Moreover, the highly correlated hand preferences between these three versions suggest that the tube task provides a more robust means of measuring manual preferences. Am J Phys Anthropol 152:315–321, 2013. © 2013 Wiley Periodicals, Inc.     

Problem-Solving Strategies and Hand Preferences for a Multicomponent Task by Tufted Capuchins (Cebus apella)

We examined hand preferences in 23 tufted capuchins (Cebus apella) in 2 tasks requiring the lid of a box to be lifted before taking out a peanut. The first task, Box 1, could entail either 2 or 3 problem-solving acts, with the 3-act solution involving bimanual coordination for food retrieval. The second task, Box 2, involved only the 3-act solution. The results indicated that the types of solution employed to perform the task influenced capuchin hand preferences. In the 2-act solution, capuchins exhibited a significant right-hand bias for the final one-handed reaching action, but not for the initial lid lifting action. In contrast, in the 3-act solution, no significant asymmetry emerged for any act. We noted a significant effect of subject's sex on the strength of laterality, with males being more strongly lateralized than females. We discuss results in the light of recent models of primate laterality.     

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.     

Hand preferences on unimanual and bimanual tasks in white-faced capuchins (Cebus capucinus)

This study examined hand preference in white-faced capuchins on a unimanual task and on a coordinated bimanual task. For the unimanual task, handedness was assessed by observing simple reaching for small grains. For the bimanual task, tubes lined with chocolate paste inside were presented to the capuchins. The hand and the finger(s) used to remove chocolate paste were recorded. Seven individuals out of eight in the reaching task and 12 out of 13 in the tube task exhibited a hand preference. Moreover, test-retest correlations showed stability in hand use across time for the coordinated bimanual task. We found no significant differences in strength of hand preference between sexes. Finally, as noted in other primate species, the capuchins were more lateralized in the bimanual task compared to the unimanual task.     

Behavioral laterality in captive bonobos (Pan paniscus): Replication and extension

We report observational data on behavioral laterality in 10 captive bonobos (Pan paniscus)at the San Diego Zoo. The unimanual measures include carrying, leading limb in locomotion, self-touching, face-touching, reaching, and gestures. We also recorded bimanual feeding in these subjects. A significant population level left-hand bias exists for carrying. Right-hand biases occur for leading limb in locomotion and gestures. During bimanual feeding, the bonobos hold food items with the left hand while feeding with the right hand. Overall, bonobos exhibit behavioral asymmetries that are similar to previous findings in other pongid ape species. The asymmetries in gestures and bimanual feeding represent novel findings with theoretical implications for the origins of tool use and language.     

Laterality in hand use across four tool-use behaviors among the wild chimpanzees of Bossou, Guinea, West Africa

Population-level right handedness is a human universal, whose evolutionary origins are the source of considerable empirical and theoretical debate. Although our closest neighbor, the chimpanzee, shows some evidence for population-level handedness in captivity, there is little evidence from the wild. Tool-use measures of hand use in chimpanzees have yielded a great deal of variation in directionality and strength in hand preference, which still remains largely unexplored and unexplained. Data on five measures of hand use across four tool-use skills--ant-dipping, algae-scooping, pestle-pounding and nut-cracking--among the wild chimpanzees of Bossou, Guinea, West Africa, are presented here. This study aims to explore age- and sex-class effects, as well as the influence of task motor, cognitive and haptic demands, on the strength and directionality of hand preference within and across all five measures of hand use. Although there was no age- or sex-class effect on the directionality of hand preference, immature 相似文献   

A study of correlates of hand preferences in squirrel monkeys (Saimiri sciureus)

The purpose of this study was to test the influence of sex, age, social rank, matriline membership, posture, and visual and tactual motor control on manual preferences inSaimiri sciureus. A well-established social group of 12 squirrel monkeys, aged 2 to 14 yrs and consisting of two matrilines with social rank known for each animal, was presented with four different food-reaching tasks and assessed for hand preferences with a minimum of 100 reaches per animal. Frequency of occurrence of hand preferences at the group level and degree of hand preferences at the individual level depended on posture and on whether the reaching act took place under visual or tactual guidance. Sex, age, social rank, and matriline membership were not found to determine frequency of occurrence, direction or degree of hand preferences with the exception of one task in which a significant negative correlation between the degree of hand preference and age was found. Nine out of 12 monkeys showed task-dependent changes in the hand they used preferentially while only three animals preferred the same hand in all four tasks. Significant preferences for the use of right or left hand on a given task were distributed almost equally between individuals. Thus, the results of this study suggest task-specific demands like posture and/or whether reaching was visually or tactually guided to be the major correlates of hand preferences in food-reaching tasks in squirrel monkeys.     

Hand preference in unimanual and bimanual tasks and postural effect on manual laterality in captive red-capped mangabeys (Cercocebus torquatus torquatus)

Hand preference in 11 captive red-capped mangabeys (Cercocebus torquatus torquatus) was examined under different conditions: a free situation during spontaneous food processing, three different postural conditions (brachiating, and bipedal and tripedal standing), and a situation involving bimanual processing. Generally, individual laterality was found regardless of the task and behavior involved. However, the number of monkeys with hand preferences and the strength of the preference increased with the complexity of the tasks. The monkeys exhibited a significantly higher and positive mean manual preference index (HI) when they were hanging than when they were quadrupedal or sitting. The strength of manual preference (ABS-HI) was in turn higher when the monkeys were hanging or bipedal than when they were quadrupedal. The strength of manual preference was higher for both the bimanual and experimental tasks than for unimanual tasks and spontaneous activities. Although our sample was too small to allow us to make any generalizations concerning lateral preferences in red-capped mangabeys, we propose some hypotheses about the influence of posture stability and task complexity.     

Hand preferences in captive gorillas,orang-utans and gibbons

Hand preference was assessed in 12 gorillas (Gorilla gorilla gorilla), 13 orang-utans (Pongo pygmaeus abelii), and 9 gibbons (Hylobates lar) by using a floor retrieval task and a mesh retrieval task. Hand preference was also assessed in 8 gorillas and 8 orang-utans by using a task involving the unfastening of a hasp. A bipedal requirement during testing (mesh retrieval task) facilitated detection of hand preferences. A significant left-hand preference was found for the gibbons with 6 of 6 gibbons preferring their left hand on the mesh retrieval task. Similarly, a significant right-hand preference was found for the gorillas with 10 of 12 gorillas preferring their right hand on the mesh retrieval task. The data for the orang-utan suggest a bimodal distribution on all tasks. Since the gibbon and gorilla in the wild engage in bipedal locomotion more frequently than the orangutan, one possible interpretation for these results correlates the degree of bipedal behavior of a species in its natural environment with its readiness to exhibit a unilateral population-level hand preference.     

Consistency of hand preference across low-level and high-level tasks in capuchin monkeys (Cebus apella)

Numerous studies investigating behavioral lateralization in capuchins have been published. Although some research groups have reported a population-level hand preference, other researchers have argued that capuchins do not show hand preference at the population level. As task complexity influences the expression of handedness in other primate species, the purpose of this study was to collect hand preference data across a variety of high- and low-level tasks to evaluate how task complexity influences the expression of hand preference in capuchins. We tested eleven captive brown capuchin monkeys (Cebus apella) to determine if they show consistent hand preferences across multiple high- and low-level tasks. Capuchins were expected to display high intertask consistency across the high-level tasks but not the low-level tasks. Although most individuals showed significant hand preferences for each task, only two of the high-level tasks that involved similar hand motions were significantly positively correlated, indicating consistency of hand preference across these tasks only. None of the tasks elicited a group-level hand preference. High-level tasks elicited a greater strength of hand preference than did low-level tasks. No sex differences were found for the direction or strength of hand preference for any task. These results contribute to the growing database of primate laterality and provide additional evidence that capuchins do not display group-level hand preferences.