The use of probing tools by tufted capuchins (cebus apella): evidence for increased right-hand preference with age

We examined hand preference in the use of tools by tufted capuchins (Cebus apella). We presented a colony of monkeys with an enclosed container designed to accommodate the use of probing tools. Over an 8-month period, 13 monkeys used probes to extract sweet syrup from the narrow opening of the apparatus. Five monkeys exhibited bias toward use of their right hand and eight monkeys exhibited bias toward use of their left hand. Adult monkeys exhibited a greater percentage of right-hand preferent probing sequences than did juveniles. These results are consistent with hypotheses that in tasks that involve the use of tools, nonhuman primates exhibit strong lateral asymmetries at the individual level, a moderate left-hand bias at the population level, and increased bias with age toward use of the right hand.     

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 preference in capuchin monkeys varies with age

The purpose of this research was to examine the influence of age on hand preference in capuchin monkeys (Cebus apella). Twenty-two capuchins, aged 6 months to 30 years, were presented with a task that involved reaching for food and a task that involved using sponging tools to absorb juice. Adults exhibited a greater percentage of right-handed actions in each task than did immature subjects. Adults also exhibited a stronger lateral bias than did immature subjects in the sponging task. These results are consistent with hypotheses: a) adult capuchin monkeys are biased toward use of their right hand for reaching; b) adult capuchins exhibit a greater incidence of right-hand preference than do immature capuchins; and c) primates exhibit age-related differences in the strength and direction of hand preference in tasks that involve the use of tools.     

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     

Hand Preferences for a Haptic Task in Chimpanzees (Pan troglodytes)

We tested the hand preferences of 20 chimpanzees (Pan troglodytes) for a haptic task requiring individuals to search for grapes in an opaque bucket filled with water. We compared these data to the hand preferences displayed by the same chimpanzees during reaching and bimanual feeding tasks. The chimpanzees displayed no significant hand preference for the reaching or bimanual feeding tasks, but exhibited a right-hand preference while performing the haptic task. In contrast, New and Old World monkeys display left-hand preferences for similar tasks. We discuss the relevance of these findings for the evolution of handedness in primates.     

Hand use by tufted capuchins (Cebus apella) to extract a small food item from a tube: digit movements, hand preference, and performance

The aim of this study was to analyze the manual patterns used by tufted capuchin monkeys (Cebus apella) to retrieve a small food item from a narrow tube, with special attention focused on the independent use of single fingers, fine digit movements, hand preference, and intermanual differences in the time it took the monkeys to obtain the food. The capuchins (n = 20) mainly used their forefinger to extract the food from the tube. The simultaneous use of the index and middle fingers occurred less frequently, and the use of the forefinger in combination with other digits occurred rarely. The capuchins demonstrated a capacity to move single digits independently when the fingers were locating the food inside the tube, and displayed a high mobility of the distal phalanx joints. However, they possessed only a limited capacity to coordinate single fingers in space, and displayed only a slight degree of manual preshaping when they approached the tube. A hand-preference analysis failed to reveal any significant lateral bias for the group, since both adults (> or = 5 years) and immature individuals (<5 years) of both sexes used either hand with the same frequency. Conversely, a latency analysis showed a significant interaction between the subject's age and performance difference between hands: in adults, but not in immature individuals, the left hand was faster than the right hand in retrieving food.     

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.     

Laterality of Hand Function in Tufted Capuchin Monkeys (Cebus apella): Comparison between Tool Use Actions and Spontaneous Non-tool Actions

This study examined hand preference for tool use and spontaneous non-tool actions in tufted capuchin monkeys (Cebus apella). We noted a lack of lateral bias across measures, and greater strength of hand preference for tool use than for self-directed or feeding activities. Animals that used tools exhibited a population-level right-hand bias for self-touching whereas animals that did not use tools exhibited a lack of lateral bias for this measure. Our findings are consistent with views that hand preference is expressed more strongly for tool use than for non-tool activities, and that lateral bias for self-directed behavior is related to problem-solving skills in primates.     

First evidence on foot preference during locomotion in Old World monkeys: a study of quadrupedal and bipedal actions in Sichuan snub-nosed monkeys (Rhinopithecus roxellana)

This is the first report of foot preference during locomotion in Old World monkeys. Foot preferences during the quadrupedal walking action and the bipedal shifting action of a naturalistic group of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in Zhouzhi National Nature Reserve in the Qinling Mountains of China were investigated. Twelve of 21 individuals tested on quadrupedal action and all 21 individuals tested on bipedal action exhibited a significant foot preference. Both significant right- and left-footed preferences were observed; sex affected neither direction nor strength of foot preference in both actions. The finding that 61.90% of focal R. roxellana showed a right-foot preference, both in quadrupedal action based on the footed index and in bipedal action based on the z-score, is in partial agreement with the postural origin hypothesis on footedness. Foot preference was significantly stronger in bipedal action than in quadrupedal action, supporting the view that posture could be a crucial factor influencing foot preference as well as hand preference in this species.     

A multi-task assessment of hand lateralization in capuchin monkeys (Cebus apella)

Eight tasks were presented to ten tufted capuchin monkeys and hand preferences were recorded for each subject on each task. The strength of hand preferences varied accross tasks. Although no significant population-level left- or right-hand bias emerged for any of the tasks, there was a tendency toward greater left-hand use in a task requiring the use of probing tools. The data also confirm that simple reaching is of limited value as a measure of hand preference. Hand preferences in capuchin monkeys appear to be determined by a number of individual, species, and environmental factors.     

Manipulative Tendencies of Captive Cebus albifrons

We conducted two experiments to examine the manipulative tendencies of captive Cebus albifrons. In Experiment 1 we examined hand preference for reaching by providing subjects with food either on the cage floor (to facilitate quadrupedal reaching) or at the height of an upright subject's shoulder (to facilitate bipedal reaching). In Experiment 2 we examined combinatorial manipulation by providing subjects with nesting containers and other portable manipulanda. Results indicate that C. albifrons exhibits greater use of the right hand for bipedal versus quadrupedal reaching (exhibiting a group-level lack of bias for bipedal reaching and a left-hand bias for quadrupedal reaching), combines objects using a simple pairing strategy, and uses and produces simple tools. Aspects of these findings parallel those for Cebus apella.     

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.     

The effect of size,age, and recent egg laying on copulatory choice of the hermaphroditic mollusc Aplysia juliana

Summary

Aplysia juliana is a cross-fertilizing, simultaneous hermaphrodite. During copulation, an individual may act as either a sperm donor or a sperm recipient, or both, when a pair copulates reciprocally. Experiments were conducted with A. juliana to determine if an animal's size, age, or recent egg-laying activity influenced its choice of copulatory role. Animals were isolated except when paired during daily, half-hour trials. In the first experiment, mature animals of different size (weight) but similar age were randomly paired. Animal size had no effect on the initial copulatory role chosen. In the second experiment, young, maturing A. juliana were paired with older animals. Young animals showed no preference in initial copulatory role either as a group or individually. Older A. juliana showed no copulatory preference as a group, but over half of the individuals demonstrated a consistent choice of one role. Some individuals acted almost exclusively as sperm donors, others as sperm recipients, suggesting that as an A. juliana ages, it is likely to develop a preference for a single copulatory role. A record of daily egg mass production was kept for all animals in the second experiment. Production of an egg mass since the last copulation did not affect the copulatory role chosen in the subsequent copulation.     

Hand preferences of captive chimpanzees (Pan troglodytes) in simple reaching for food

We examined chimpanzee hand preference in simple reaching for food, with special reference to manipulative patterns and the developmental shift. We observed 80 captive chimpanzees, ranging from 1 to 25 years old. We also studied the manipulative patterns (grip- types) of 70 individuals as they reached for raisins scattered randomly on the floor. We employed LQ score as a measure of hand preference and designated the subjects right- handers (or left- handers) if they used their right hands (left hands) above chance level. Although the numbers of right- handers and left- handers are almost equal, the distribution of the strength is not symmetrical in both groups. Strong preference was exhibited by more left- handers than right- handers. Subjects > 9 years old exhibited greater hand preference, whereas subjects < 9 years old were ambidextrous. We classified manipulative patterns for reaching into five basic grip- types and analyzed them vis- à- vis age. There is no significant correlation between preferred hand and manipulative patterns. However, adult subjects tended to use an index- and - middle- finger grip with the left hand and to use imprecise grips with the right hand more often than other patterns regardless which hand they preferred. These data demonstrate a developmental shift in hand preference and manipulative patterns and also reveal functional asymmetries between the right and the left hand in Pan troglodytes.     

Black lemur (Lemur macaco) hand preference in food reaching

The present study assessed the hand preferences exhibited by 33 black lemurs during routine feeding. Individual animals displayed hand preferences that were consistent across observations separated by as much as seven months. Within this population, 20 were left hand preferent, 12 right preferent, and 1 was ambidextrous. Correlational analysis of age and percentage left hand use indicated an inverse relationship in which younger animals tended towards the preferential use of the left hand and older animals the preferential use of the right hand. Similar analysis found no relationship between either sex and hand preference or familial relationship and hand preference. The skewed distribution of age in this sample renders tentative conclusions regarding age-related variations in hand preference. It is suggested that if the hand preferences of the black lemur are not age-related, then this species may be characterized as having a bias towards the preferential use of the left hand for food reaching.     

Tube task hand preference in captive hylobatids

The link between laterality in humans and other primates is still hotly debated. Hylobatids have been rather neglected in this research area, yet they can provide important insights because: (1) they share with humans a complex vocal repertoire, which in humans is thought to be associated with brain hemispheric specialization and lateralized behaviors; (2) their adaptation to arboreality has produced unique postural constraints; (3) the little that is known about laterality in gibbons is contradictory (captive studies have provided conflicting results, while a field study on siamangs reported a population-level left-hand preference). To clarify this, we investigated hand preference in captive hylobatids [n = 42; 22 siamangs (Symphalangus syndactylus) and 20 gibbons (Hylobates sp., Nomascus leucogenys)] in nine Japanese facilities. We had a large sample size, controlled for possible confounds (posture, enclosure limitations) and used a well-established testing protocol (tube task). Handedness indices calculated from raw frequencies and bouts were highly correlated and showed a significant left-hand skew, which is consistent with data from wild siamangs. Major differences between captive and wild siamangs were a larger number of ambiguously handed individuals, and no significant age-related variation in captivity. The use of the index finger elicited a much more strongly lateralized response than the thumb. These results confirmed a left-hand preference in siamangs, but were equivocal in other hylobatids, and suggest selective pressures that may have acted on the highly arboreal hylobatids to favor handedness. Our study also indicates factors that might explain the discrepancy in the literature between handedness studies on captive and wild primate populations.     

Hand Preference in Free-Ranging White-Throated Capuchins (Cebus capucinus) in Costa Rica

I studied the hand preference patterns of individuals in three troops of white-throated capuchins (C. capucinus) in Palo Verde, Costa Rica, during 11 months from February 1995 to January 1996. I used focal and ad libitum sampling techniques and tested several frameworks that seek to explain and to predict primate hand use patterns via a variety of spontaneous tasks that differ in manipulative difficulty and required postural regulation: reach, tap, grab, carry, and object-use. The monkeys showed symmetrical hand use patterns for the easy tasks, reach and tap; strongly asymmetrical patterns for the highly manipulative task, object-use, at the individual level; and weak population-level biases for tasks requiring a degree of postural regulation, carry. The results for data on grab are inconclusive. These results do not support the available primate hand use frameworks and differ from most of the captive literature on hand preference in Cebus. The findings indicate that postural regulation may influence hand use patterns in nonhuman primates at the population level.     

A Comparative Assessment of Hand Preference in Captive Red Howler Monkeys,Alouatta seniculus and Yellow-Breasted Capuchin Monkeys,Sapajus xanthosternos

There are two major theories that attempt to explain hand preference in non-human primates–the ‘task complexity’ theory and the ‘postural origins’ theory. In the present study, we proposed a third hypothesis to explain the evolutionary origin of hand preference in non-human primates, stating that it could have evolved owing to structural and functional adaptations to feeding, which we refer to as the ‘niche structure’ hypothesis. We attempted to explore this hypothesis by comparing hand preference across species that differ in the feeding ecology and niche structure: red howler monkeys, Alouatta seniculus and yellow-breasted capuchin monkeys, Sapajus xanthosternos. The red howler monkeys used the mouth to obtain food more frequently than the yellow-breasted capuchin monkeys. The red howler monkeys almost never reached for food presented on the opposite side of a wire mesh or inside a portable container, whereas the yellow-breasted capuchin monkeys reached for food presented in all four spatial arrangements (scattered, on the opposite side of a wire mesh, inside a suspended container, and inside a portable container). In contrast to the red howler monkeys that almost never acquired bipedal and clinging posture, the yellow-breasted capuchin monkeys acquired all five body postures (sitting, bipedal, tripedal, clinging, and hanging). Although there was no difference between the proportion of the red howler monkeys and the yellow-breasted capuchin monkeys that preferentially used one hand, the yellow-breasted capuchin monkeys exhibited an overall weaker hand preference than the red howler monkeys. Differences in hand preference diminished with the increasing complexity of the reaching-for-food tasks, i.e., the relatively more complex tasks were perceived as equally complex by both the red howler monkeys and the yellow-breasted capuchin monkeys. These findings suggest that species-specific differences in feeding ecology and niche structure can influence the perception of the complexity of the task and, consequently, hand preference.     

Locomotor behavior,body size,and comparative ecology of seven Surinam monkeys

The locomotor behavior, of seven sympatric species of New World monkeys—Saguinus midas midas, Saimiri sciureus, Pithecia pithecia, Chiropotes satanas chiropotes, Cebus apella apella, Alouatta seniculus, and Ateles paniscus panisus—was studied at the Raleighvallen-Voltzberg Nature Reserve in Central Surinam. This paper examines the way in which locomotor behavior is related to body size and to ecological parameters such as forest stratification, forest type, and diet. Locomotor behavior is clearly related to the size of the species; with increasing size, the amount of climbing increases and the amount of leaping decreases. In general, larger monkeys use larger arboreal supports; however, Saguinus midas midas uses relatively larger, and Ateles paniscus paniscus relatively smaller supports than expected from body size alone. Leaping is associated with use of the forest understory and with use of liane forest. Other types of locomotion are associated with main canopy use in a variety of forest types. There are no consistent associations between diet and either locomotor behavior or forest utilization; rather, monkeys with similar diets show locomotor and habitat differentiation.     

Handedness found in a wild group of moor monkeys Macaca maurus in the Karaenta Nature Reserve, South Sulawesi,Indonesia

Hand preference of wild moor monkeys Macaca maurus was investigated in food reaching situations at the Karaenta Nature Reserve, South Sulawesi, Indonesia. The frequency picking up sweet-corn grains to take into the mouth by either hand of monkeys was counted directly at the feeding ground where sweet-corn grains were scattered for monkeys. Among the 20 monkeys examined, 8 were fight-handed, 8 were left-handed, and 4 were ambilateral. The results indicated the prevailed hand preference on individual level but not either trends of left- or right-hand preference on population level. The trend toward a higher proportion of left-handed monkeys found in Japanese and rhesus monkeys was not found. Some other characteristics found in moor monkeys are discussed in comparison with those previous findings in Japanese and Tibetan macaques in order to evaluate variations within the genus Macaca.