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   

Clapping in chimpanzees: evidence of exclusive hand preference in a spontaneous, bimanual gesture

An increased understanding of spontaneous bimanual tasks performed by chimpanzees would enhance the ongoing debate on population handedness in this species, and thus shed light on the evolution of hemispheric lateralization in humans. However, documentation of such bimanual activities has been largely absent in the literature because such behavior is infrequently observed in the natural repertoire of this species. This study presents data on a simple, spontaneous bimanual gesture-"clap"-that was investigated in a naturalistic group of 26 chimpanzees (Pan troglodytes). Sixteen individuals exhibited a total of 657 bouts of clapping directed toward humans, usually in the context of food availability. Most individuals were exclusive in their dominant hand preference, but since there was no population bias to the right or the left, the population is placed at level 3 of McGrew and Marchant's [Yearbook of Physical Anthropology 40:201-232, 1997] laterality framework. This is the first reported evidence of level 3 laterality in a non-tool-using task. Clapping increased in frequency with age, being common in adults, present at lower rates in adolescents, and absent in infants and juveniles. There was no effect of rearing or sex. The lack of population bias to the left or right for this bimanual gesture has implications for the debate on the evolution of language.     

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.     

Laterality in spontaneous motor activity of chimpanzees and squirrel monkeys

Varieties of nonmanipulative motor responses were observed in chimpanzees and squirrel monkeys. Chimpanzees displayed a right hand preference for touching their inanimate environments but used their right and left hands equally for touching their faces and their bodies. The latter result was not consistent with previous reports of a left hand preference for face touching in apes. The right hand preference for environmental touching was stronger in male than in female chimpanzees. Squirrel monkeys had a right preference for combined hand and foot responses directed to their bodies, but expressed no handedness for environmentally directed touching. These limb preferences in chimpanzees and squirrel monkeys indicate that neither precise, complex manipulation nor postural instability are necessary conditions for population level hand preferences. Factor analysis of the chimpanzee manual responses showed distinct self and environmentally directed factors. Analysis of the squirrel monkey data also showed self and environmental factors, except that body scratching had a negative loading on the environmental factor. This latter result suggests that self-scratching by squirrel monkeys is a displacement activity that suppresses manual exploration of the environment. © 1992 Wiley-Liss, Inc.     

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.     

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.     

Simple Reaching Is Not So Simple: Association Between Hand Use and Grip Preferences in Captive Chimpanzees

We assessed the relationship between grip preference and hand use in chimpanzees in 2 experiments. In experiment 1, we evaluated consistency in hand use and grip preference across 4 food types. The chimpanzees showed population-level right-handedness and there are significant positive associations for both hand and grip use across food types. In experiment 2, we assessed validity of hand use in relation to grip preference in 2 colonies of chimpanzees via the same methodology. Differences in hand preferences between colonies were associated with variation in the observed grip preferences. There was no evidence of rearing effects on handedness in either colony. We discuss the overall results in the context of the evolution of handedness in relation to increasing motor demands as manifest in variation on grasping behavior.     

Bipedal tool use strengthens chimpanzee hand preferences

The degree to which non-human primate behavior is lateralized, at either individual or population levels, remains controversial. We investigated the relationship between hand preference and posture during tool use in chimpanzees (Pan troglodytes) during bipedal tool use. We experimentally induced tool use in a supported bipedal posture, an unsupported bipedal posture, and a seated posture. Neither bipedal tool use nor these supported conditions have been previously evaluated in apes. The hypotheses tested were 1) bipedal posture will increase the strength of hand preference, and 2) a bipedal stance, without the use of one hand for support, will elicit a right hand preference. Results supported the first, but not the second hypothesis: bipedalism induced the subjects to become more lateralized, but not in any particular direction. Instead, it appears that subtle pre-existing lateral biases, to either the right or left, were emphasized with increasing postural demands. This result has interesting implications for theories of the evolution of tool use and bipedalism, as the combination of bipedalism and tool use may have helped drive extreme lateralization in modern humans, but cannot alone account for the preponderance of right-handedness.     

Tool-using and -making behavior in wild chimpanzees at Bossou,Guinea

The behavior of wild chimpanzees at Bossou, Guinea, was studied from November 1976 to May 1977 recognizing each chimpanzee without artificial feeding. During the study period some tool-using and tool-making behavior was observed, as follows: (1) Although water drinking using a “leaf-sponge” was not seen, that using a “leaf-spoon” was observed for taking water from the hollow of a tree. (2) “Termite fishing” was not seen in this group although there were many termite hills in the moving range of the chimpanzees. They dug termites from the hollow of a tree by pounding with a small stick. Similar use of a stick was made for digging up the resin from a tree. (3) “Aimed throwing” was frequently observed in adult males for attacking an observer, and in adolescents and juveniles as mischief against an observer or for their own play. (4) “Nut cracking” with a pair of stones was seen for removing the ovule from palm-seeds. Particular stones were repeatedly used by many chimpanzees for a long period. (5) “Branch hauling” represented difficult work. Patient and inventive manufacture of proper sticks was necessary for capturing branches which they were unable to reach normally. Local variations in the tool-using patterns and manufacturing ability of chimpanzees are discussed.     

Limb use and preferences in wild orang-utans during feeding and locomotor behavior

Limited data are available on hemispheric lateralization in wild orang-utans. There has been only one previous investigation of limb preferences in wild orang-utans [Yeager, 1991]. We examined the lateralization of limb use in wild Bornean orang-utans (Pongo pygmaeus pygmaeus) with the aim of providing more insight into possible hemispheric specialization in wild nonhuman primates. Here, we report in detail on limb use and preference during arboreal locomotion between trees (N=6) and on feeding involving one limb (N=8) and two limbs (N=6). We distinguished between locomotion between overlapping trees (Type I) and locomotion involving gap crossing (Types II and III). For locomotion Type I, the six orang-utans showed no leading hand preference, however for locomotion Types II and III, all six showed significant right-hand preferences. All eight orang-utans showed individual hand preferences for reaching for food, but no significant group bias was found. Limb preferences for feeding involving two limbs (hand-hand or hand-foot) differed between juveniles (right hand-right foot), adult females (left hand-right hand) and adult males (right hand-left hand). Although not present for all tasks, the results indicate that orang-utans do show evidence of hemispheric specialization, but the use of the hands is not under a strong lateralized hemispheric control and is adaptable.     

Propagation of handclasp grooming among captive chimpanzees

A grooming posture previously reported for two wild chimpanzee (Pan troglodytes) communities developed spontaneously in a captive group of the same species. This offered a unique opportunity to follow the propagation of a new social custom. The posture consists of two partners grasping hands—either both right hands or both left hands—and raising the arms in an A-frame above their heads while mutually grooming with their free hands. The propagation of this pattern was followed over a 5 year period. In the beginning, handclasps were always initiated by the same adult female. This female initiated the posture mainly with her adult female kin. In subsequent years, these relatives became frequent participants in the posture with each other as well as with nonrelatives. Over the years the posture increased in frequency and duration and spread to the majority of adults and also to a few adolescents and older juveniles. The pattern persisted after removal of the apparent originator. Am. J. Primatol. 43:339–346, 1997. © 1997 Wiley-Liss, Inc.     

Relationship between physiological tremor and cognitive function in physically active older women

[Purpose]This study aimed to compare the physiological tremor, grip strength, and cognitive function of sedentary and physically active older adults.[Methods]Twenty-four older adults aged ≥65 years participated in this study and were divided into the sedentary (76.5±4.4 years, n=12) and physically active (73.5±3.3 years, n=12) groups. Each group completed the Mini-Mental State Examination (MMSE) for cognitive function assessment. Physiological tremor was measured using an accelerometer for both hands at rest and the left/right hand with a 1,000 g dumbbell on the palm in neutral positions and the elbow flexed at 90°. Physical fitness was measured by grip strength and completion of the Short Physical Performance Battery (SPPB) and the 6-min walk test.[Results]The physically active group showed a significantly lower level of physiological tremor in both hands at rest and the left/right hand with a 1,000 g dumbbell on the palm (P<0.05) than that in the sedentary group. For cognitive function, the physically active group showed significantly higher scores than those in the sedentary group (P<0.001). No significant correlation was found between cognitive function and left/right grip strength (left: r = 0.117, P = 0.585; right: r = 0.230, P = 0.279), physiological tremor in both hands at rest (left: r = -0.524, P < 0.001; right: r = -0.508, P < 0.05), and the left/right hand with a 1,000 g dumbbell on the palm (left: r = -0.505, P < 0.05; right: r = -0.458, P < 0.05).[Conclusion]Physiological tremor of the hands has the potential to be a useful predictor of cognitive function in older adults.     

Age-related differences in corticospinal control during functional isometric contractions in left and right hands.

The purpose of the study was to examine age-related differences in electromyographic (EMG) responses to transcranial magnetic stimulation (TMS) during functional isometric contractions in left and right hands. EMG responses were recorded from the first dorsal interosseus muscle following TMS in 10 young (26.6 +/- 1.3 yr) and 10 old (67.6 +/- 2.3 yr) right-handed subjects. Muscle evoked potentials (MEPs) and silent-period durations were obtained in the left and right hands during index finger abduction, a precision grip, a power grip, and a scissor grip, while EMG was held constant at 5% of maximum. For all tasks, MEP area was 30% (P < 0.001) lower in the left hand of old compared with young subjects, whereas there was no age difference in the right hand. The duration of the EMG silent period was 14% (P < 0.001) shorter in old (150.3 +/- 2.9 ms) compared with young (173.9 +/- 3.0 ms) subjects, and the age differences were accentuated in the left hand (19% shorter, P < 0.001). For all subjects, the largest MEP area (10-12% larger) and longest EMG silent period (8-19 ms longer) were observed for the scissor grip compared with the other three tasks, and the largest task-dependent change in these variables was observed in the right hand of older adults. These differences in corticospinal control in the left and right hands of older adults may reflect neural adaptations that occur throughout a lifetime of preferential hand use for skilled (dominant) and unskilled (nondominant) motor tasks.     

Grip preference, dermatoglyphics, and hand use in captive chimpanzees (Pan troglodytes)

This paper examined the association between grip type, hand use, and fingerprint patterns in a sample of captive chimpanzees. Grip type for simple reaching was assessed for the left and right hand and classified as thumb-index, middle-index, or single-digit responses. Fingerprint patterns were characterized as whorls, loops, or arches on each finger. The results indicated that chimpanzees exhibit significantly more thumb-index responses for the right compared to the left hand. In addition, thumb-index responses were more prevalent for subjects that had a whorl compared to a loop or arch on their thumb. The results suggest that fingerprint patterns are associated with individual differences in grasping type in chimpanzees as well as some variation in hand use.     

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.     

Spatial compatibility and position of the effectors

Mean reaction times obtained with crossed hands (right had on the left and left hand on the right) are slower than reaction times obtained with uncrossed hands (right hand on the right and left hand on the left). These results have been explained as a compatibility effect between the responding hand and its spatial position. The goal of the present experiment was to establish whether the position of the hand is encoded by the subjects relative to their body (absolute position) or relative to the other hand (relative position). The subjects performed a discrimination task on two visual stimuli. Stimuli and hands were either on the same side of the body (both on the left or both on the right) or had different absolute position. In all conditions the subjects responded with crossed and uncrossed hands. The results support the hypothesis that relative position is encoded.     

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 abnormalities in pygmy chimpanzees (Pan paniscus)

Hand deformities in 2 wild-born pygmy chimpanzees were investigated through dissection, linear measurements, bone mineral analysis and X-rays. Weight of tissue (skin, muscle, bone) was determined for each body segment (hand, forearm, upper arm), permitting comparison between the abnormal and normal upper limbs. In one individual, the right and left forearms and hands differed in tissue weights and bone mineral content. In the other, the hands differed in tissue composition, but not the forearms.     

Adult and juvenile bearded capuchin monkeys handle stone hammers differently during nut-cracking

Wild bearded capuchin monkeys (Sapajus libidinosus) habitually use stone hammers to crack open palm nuts and seeds on anvils. This activity requires strength, balance, and precise movement of a large stone with respect to the item placed on an anvil. We explored how well young monkeys cope with these challenges by examining their behavior and the behavior of adults while they cracked palm nuts using a stone. Using video records, we compared actions of six juvenile (2–5 years) and six adult (7+ years) wild monkeys during their first 20 strikes with one unfamiliar ellipsoid, quartzite stone (540 g), and the outcomes of these strikes. Compared with adults, juveniles cracked fewer nuts, performed a more diverse set of exploratory actions, and less frequently placed one or both hands on top of the stone on the downward motion. Adults and juveniles displayed similar low frequencies of striking with a slanted trajectory, missing the nut, and losing control over the nut or stone after striking. These findings indicate that young monkeys control the trajectory of a stone adequately but that is not sufficient to crack nuts as effectively as adults do. Compared with juveniles, adults more quickly perceive how to grip the stone efficiently, and they are able to adjust their grip dynamically during the strike. Young monkeys develop expertise in the latter aspects of cracking nuts over the course of several years of regular practice, indicating that perceptual learning about these aspects of percussion occurs slowly. Juvenile and adult humans learning to use stones to crack nuts also master these features of cracking nuts very slowly.