A report of behavioral lateralization in an infant orang-utan (Pongo pygmaeus)

The limb used to perform seven common activities was recorded during weekly observations of an infant orang-utan. For five of these behaviors, preferences were found that remained consistent in their direction, although there were week to week fluctuations in magnitude. Most notably, a right hand preference was found for nonfood reaching and a right hindlimb preference appeared for initiating locomotion. Although initially, food reaching was predominantly with the right hand, a shift toward preferential left hand use occurred during the final weeks of the study. Additionally, a left hand preference appeared when the infant touched either its body or head.     

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.     

Laterality in semi‐free‐ranging black and white ruffed lemurs (Varecia variegata variegata): head‐tilt correlates with hand use during feeding

Previous studies in human and chimpanzee infants have identified a predictive relationship between early rightward head orientation and later right hand use. Data from lemurs suggest a leftward bias in hand preference, but there are no data on head positioning. The purpose of this study was to examine the relationship between head and hand preferences in the black and white ruffed lemur (Varecia variegata variegata). Ruffed lemurs rotate the head vertically during chewing in a behavior called head‐tilting. Frequency of head‐tilting and bouts of unimanual hand use were measured during normal feeding in a semi‐free‐ranging population of lemurs. Subjects were provisioned at feeding platforms twice daily with fresh fruits, vegetables, and other food items. Sampling was spontaneous and all observations were videotaped. No group‐level bias was found for head‐tilting, but a left hand bias emerged for hand use. A positive relationship was found between direction of head‐tilting preference and direction of hand use preference such that left head‐tilts increased as left hand use increased. Furthermore, left head‐tilts increased as the degree of hand preference lateralization increased. When the hand used to bring food to the mouth just before head‐tilting was examined, there was a strong bias for the left hand to precede left head‐tilts. For right head‐tilts, however, lemurs were equally likely to use either hand before head‐tilting. Overall a strong relationship was found between the left hand and left head‐tilting in black and white ruffed lemurs, suggesting a common link between these behaviors. However, the direction of bias was different from that seen in human and chimpanzee studies. Additional studies on patterns of laterality would be informative for understanding how laterality has changed across the primate order and the adaptive significance of laterality in primates. Am. J. Primatol. 71:1032–1040, 2009. © 2009 Wiley‐Liss, Inc.     

Laterality in quadrupedal and bipedal prosimians: Reach and whole-body turn in the mouse lemur (Microcebus murinus) and the galago (Galago moholi)

Lateral preferences in food reaching and whole-body turning were assessed in 24 prosimian primates: 16 galagos (Galago moholi) and 8 mouse lemurs (Microcebus murinus). Reach and turn preference in the galago were examined using a single testing procedure; separate procedures were used to evaluate reach and turn preferences in the mouse lemur. The motoric requirements for both species were 1) reach into a Plexiglas container with a single hand for a food item and 2) execution of a whole-body rotation about the long axis of the body. Twenty-three of the subjects tested had a reach preference (15 right, 8 left preferent) and 22 a turn preference (1 right and 21 left preferent). No correlation between the direction of reach and turn preference was found for either species; however an association between right reach preference and female gender was found for the galagos. A species difference in the strength of lateralization was found, with the more bipedal species, the galago, more strongly lateralized in both measures. This result supports the hypothesis that the assumption of upright posture was an important factor in the phylogenetic development of primate lateralization.     

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.     

Lateral bias in feeding and brachiation inHylobates

Lateralized hand use in gibbons was assessed for both food reaching and leading limb in brachiation. Sex and age effects were found in hand preference for food reaching. Adult females were all very strongly right hand preferent, whereas adult males had no across group consistent preference. Within the female group there was a strong correlation between age and strength of right handedness. When compared in terms of absolute strength of hand preference, females were found to be more strongly lateralized than males. Leading limb preference in brachiation was scored into vocal and non-vocal categories. Three subjects had a shift in preferred leading limb from the non-vocal brachiation condition to the vocal brachiation condition. This shift may be influenced by the arousal effects of species typical vocalization. The results of this study underline the importance of consideration of such factors as sex and age when interpreting behavioral lateralization data. The exploration of laterality in many different response measures is important to the achievement of a complete understanding of behavioral lateralization in 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 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.     

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.     

Baseline cortisol levels and social behavior differ as a function of handedness in marmosets (Callithrix jacchus)

Population hand preferences are rare in nonhuman primates, but individual hand preferences are consistent over a lifetime and considered to reflect an individual's preference to use a particular hemisphere when engaged in a specific task. Previous findings in marmosets have indicated that left‐handed individuals tend to be more fearful than their right‐handed counterparts. Based on these findings, we tested the hypotheses that left‐handed marmosets are (a) more reactive to a social stressor and (b) are slower than right‐handed marmosets in acquiring a reversal learning task. We examined the hand preference of 27 male and female marmosets (ages of 4–7 years old) previously tested in a social separation task and a reversal learning task. Hand preference was determined via a simple reaching task. In the social separation task, monkeys were separated from their partner and the colony for a single 7‐hr session. Urinary cortisol levels and behavior were assessed at baseline, during the separation and 24 hr postseparation. Hand preferences were equally distributed between left (n = 10), right‐handed (n = 10), and ambidextrous (n = 7) individuals. The separation phase was associated with an increase in cortisol levels and behavioral changes that were similar across handedness groups. However, cortisol levels at baseline were positively correlated with right‐handedness, and this relationship was stronger in females than in males. In addition, the occurrence of social behaviors (pre‐ and postseparation) was positively correlated with right‐handedness in both sexes. Baseline cortisol levels did not correlate significantly with social behavior. Acquisition of the reversals was poorer in females than males but did not differ as a function of handedness. We conclude that (a) both stress reactivity and cognitive flexibility are similar across handedness groups and (b) left‐handers exhibit less social behavior and have lower basal cortisol levels than ambidextrous and right‐handed subjects. The underlying causes for these differences remain to be established.     

Hand preferences in captive orangutans (Pongo pygmaeus)

The strength of the evidence for population-level handedness in the great apes is a topic of considerable debate, yet there have been few studies of handedness in orangutans. We conducted a study of manual lateralization in a captive group of eight orangutans (Pongo pygmaeus) ranking the degrees of manual preference according to a defined framework. We analyzed five behavioral patterns: eat (one- and two-handed), make/modify tool, oral tool-use, and manual tool-use. Although some individuals showed significant manual preferences for one or more tasks, at the group-level both one-handed and two-handed eating, oral tool-use, and make/modify tool were ranked at level 1 (unlateralized). Manual tool-use was ranked at level 2, with four subjects demonstrating significant hand preferences, but no group-level bias to the right or left. Four subjects also showed hand specialization to the right or left across several tasks. These results are consistent with most previous studies of manual preference in orangutans. The emergence of manual lateralization in orangutans may relate to more complex manipulative tasks. We hypothesize that more challenging manual tasks elicit stronger hand preferences.     

True handedness in the cotton-top tamarin (Saguinus oedipus)?

Hand use for 8 activities was studied in 20 captive cotton-top tamarins (Saguinus oedipus). The purpose of the study was to discover if hand preferences existed across tasks and across subjects, and, if so, whether these followed the patterns of preference inMacNeilage et al.'s (1987) “postural origins” theory of the evolution of hand preference and hemispheric specialization in primates. This theory suggests that, for haplorines, the right hand is used preferentially for manipulative acts while the left hand is used preferentially for visually guided acts. The study showed statistically significant right hand preferences for six of the seven actions which produced sufficient data to be tested. The strength and consistency of this preference makes this study the first to suggest true handedness in a non-human primate species. Deviations from the pattern were seen in scratching and other actions when performed in a vertical posture, but these deviations took the form of a weakening of the right hand preference rather than a reversal. The findings of the study may be seen as supportingMacNeilage et al.'s (1987) theoryonly if the actions studied are all considered to be manipulative, which can be argued to be the case, though some of the actions were also visually guided.     

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.     

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.     

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.     

Age-related changes in hand grip strength among rural and urban Haryanvi Jat females

The aim of this study was to investigate the age-related changes in hand grip strength (both right and left hand) among rural and urban Jat females and its correlation with various anthropometric variables. The present cross-sectional study was based on a sample of 600 Jat females (rural=300, urban=300), ranging in age from 40 to 70 years. Findings indicated that rural and urban Jat females exhibit an age-related decline in the mean values of hand grip strength (both right and left hand). The comparison suggested that rural Jat females had significantly higher overall mean value of hand grip strength (kg) (right hand=20.35 vs 18.87, p<0.001 and left hand=17.05 vs 15.68, p<0.001) than their urban counterparts. Mean values for right hand grip strength were found to be higher than left hand grip strength in rural as well as urban Jat females. Significant and positive correlation coefficients exist between hand grip strength (both right and left hand) and various anthropometric variables (height, weight, body mass index, biceps and triceps skinfold thickness) in both rural and urban Jat females. An age-related decline in manual functioning of rural and urban Jat females may be attributed to behavioral factors (reduced physical activity, sedentary lifestyle) and poor nutrition (deteriorating quality and quantity of food intake with increasing age) associated with the normal aging process.     

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.     

Patterns and laterality of hand use in free-ranging aye-ayes (<Emphasis Type="Italic">Daubentonia madagascariensis</Emphasis>) and a comparison with captive studies

We observed hand use in free-ranging aye-ayes (Daubentonia madagascariensis) on an island in the Mananara River, eastern Madagascar. The results were compared with those of two conflicting studies on hand laterality in captive aye-ayes. We argue that patterns of hand preference in wild aye-ayes are comparable to those of captive animals and that discrepancies between studies are—at least partly—caused by different ways of collecting and processing data. Aye-ayes fit Level 2 of the categories of hand laterality described by McGrew and Marchant (Yearb Phys Anthropol 40:201–232, 1997), with some individuals showing significant hand preference, but with the proportion of right- to left-preferent animals being very close to 1:1. We observed hand preference to be consistent for two of the most frequent behaviors, tapping and probing with fingers. Reaching and holding objects in hands is rare in aye-ayes, and the patterns of hand use in aye-ayes are therefore not directly comparable with those of other prosimians in which laterality has been studied. We detected no effect of sex on hand preference and were unable to determine whether there is an effect of age. The posture adopted by the animals did not influence hand preference.     

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.     

Hand preference study in marmosets (Callithrix jacchus) using food reaching tests

Hand preference has been investigated in New World primates but the data obtained thus far are controversial. In this study we investigated hand preference in common marmosets,Callithrix jacchus, during the execution of a reaching for food task. We used 46 adult common marmoset males (n=27) and females (n=19) from the Universidade of Rio Grande do Norte colony, both wild and captive-born. To test the hand preference we used a device measuring 10 cm², with a central hole 1 cm in diameter, to force the animal to use only one hand to reach for food on a food dish located underneath. Each animal was tested 5 times and had to make a maximum of 20 successful attempts per session. A total of 100 successful attempts per animal and 4,600 successful attempts for all animals were recorded during the experiment. Latency and duration of the sessions were measured and we found preference for the use of one of the hands in common marmoset individuals, i.e. 45 of total of 46 animals used significantly more the right or the left hand when performing the task. However no bias at the population level was found. Females born in captivity presented an increase in the duration of latency for the first successful attempt and in the total duration of the test sessions. These findings might be indicating differences associated with a natural tendency for females to be more selective and to spend more time exploring alimentary sources. Additionally, captive-born females may have a constrain in developing cognitive abilities regarding foraging since they have food available during most part of the time.