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.     

Patterns of hand and mouth lateral biases in bamboo leaf shoot feeding and simple food reaching in the gentle lemur (Hapalemur griseus)

Lateralized feeding behaviors of 7 wild-born and 6 captive-born Hapalemur griseus sp. were evaluated in two conditions, simple food reaching and species-typical bamboo foraging. Gentle lemurs were found to have the strongest lateral hand preference in simple food reaching of any prosimian species yet studied. However in this small sample, population-level lateralization was not found: 5 preferred the right hand and 8 the left. The movement pattern used by hapalemurs when feeding on bamboo leaf shoots was categorized into four components: PULL IN, COUNTERFORCE, TURN, and FEED IN. Lateral hand bias was scored for all four components and lateral mouth preference was scored for the two components that involved the mouth (COUNTERFORCE and FEED IN). Strong positive correlations were found between hand preference in the most lateralized components of shoot eating (TURN and FEED IN) and simple food reaching. This suggests that the gentle lemurs' species-typical bamboo feeding behavior may contribute to the exceptionally strong hand preference for simple food reaching seen in this species. There were negative correlations between the component measure of shoot feeding that required strength (COUNTERFORCE) and the components that required manipulation (TURN and FEED IN). Within the COUNTERFORCE and FEED IN components, mouth preference and hand preference were positively associated in a manner that suggested one side of the mouth was preferred for removing the shoot and the other side for mastication. Asymmetrical dental wear may provide a means of dating behavioral lateralization in ancestral species. © 1993 Wiley-Liss, Inc.     

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.     

川金丝猴摄食过程中“手”的选择使用

本文对20只川金丝猴(Rhinopithecus roxellana) 摄食过程中左右“手”的使用频次进行了观察和分析。在样本较小的条件下, 尽管被研究动物中有一半个体明显地倾向于使用右手, 但没有发现统计学上明显的种群水平的单手利现象。虽然本研究中成体金丝猴使用右手的比例在数量上高于幼体, 但在统计学上, 单手利行为不存在明显的年龄差别; 同样, 单手利用行为也不存在统计学上明显的性别差异。建议在未来的研究中采用大样本在解决复杂问题过程中对单手利行为做进一步的探讨。     

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     

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.     

Formation of lateralized motor skills in rats

Adult Wistar rats were trained to get food with a forepaw from a narrow tube under conditions of free choice of a limb. It was shown that the presence or absence of limb preference determined by preliminary short-run testing far from always defines the character and extent of lateralization formed in the process of training. It was also found that reaching the maximal extent of lateralization requires, on average, more extended period of training for the animals, which finally constituted the group of right-handers, than for the rats finally classified as left-handers.     

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 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 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.     

Manual lateralization in early primates: a comparison of two mouse lemur species

In humans, 90% of the population is right handed. Although population hand preference has been found in some primate species, the evolution of manual lateralization in primates is not yet clear. To gain insight into manual lateralization of ancestral primates, we studied hand usage in unspecialized quadrupedal, nocturnal lemurs, using a large sample size. We compared two closely related mouse lemur species to explore the variation of hand preference within the same genus. We tested 44 gray mouse lemurs and 19 Goodman's mouse lemurs in a forced food grasping task. The tests were videotaped. Measures of hand preference (i.e. the hand that is spontaneously chosen for a specific task) and successful hand usage (i.e. the hand that is successful in completing a specific task) were taken to explore manual lateralization. Both species showed manual lateralization at an individual, but not at a population level. Goodman's mouse lemurs showed stronger individual hand preferences than gray mouse lemurs. This suggests that strength in hand preference is variable within the same genus. No sex and age effects were found. The hand preference of offspring was negatively correlated to that of their mothers, but not correlated to that of their fathers. Thus, no clear genetic effect can be derived from these results. In the Goodman's mouse lemurs, hand preference increased with increasing task experience. However, successful hand usage was not affected by task experience, suggesting that successful hand usage is a more stable measurement for manual lateralization than hand preference.     

Lateralization of Motor Functions in Early Human Ontogeny

This study was aimed at testing the hypothesis about the development of lateralization of frontal motor control (FMC) as a source of preference of the right hand human in early ontogeny. The sample consisted of 108 children aged from 8 months (the initial stage of maturation of inhibitory control in reaching) to 11 months (the definitive stage). The main results showed the following: (1) at the population level, the preference of the right hand in the performance of tests that require a definite level of development of FMC appears only at the age of 11 months, i.e., synchronously with development of frontal inhibitory control of reaching; (2) the preference of the right hand is substantially stronger in 11-month-old children with a higher level of development of inhibitory motor control as compared to other children of the same age; (3) the preference of the right hand is most evident for those movements that require inhibitory control (i.e., controlled reaching). On the whole, the data obtained show that the appearance of a preference for the right hand in the second half-year of the life of an infant is caused by the development of cortical inhibitory control over motor acts (FMC), associated with the maturation of the frontal neocortex.     

Preference for conspecifics evolves earlier in males than females in a sexually dimorphic radiation of fishes

Speciation by sexual selection is generally modeled as the coevolution of female preferences and elaborate male ornaments leading to behavioral (sexual) reproductive isolation. One prediction of these models is that female preference for conspecific males should evolve earlier than male preference for conspecific females in sexually dimorphic species with male ornaments. We tested that prediction in darters, a diverse group of freshwater fishes with sexually dimorphic ornamentation. Focusing on the earliest stages of divergence, we tested preference for conspecific mates in males and females of seven closely related species pairs. Contrary to expectation, male preference for conspecific females was significantly greater than female preference for conspecific males. Males in four of the 14 species significantly preferred conspecific females; whereas, females in no species significantly preferred conspecific males. Relationships between the strength of preference for conspecifics and genetic distance revealed no difference in slope between males and females, but a significant difference in intercept, also suggesting that male preference evolves earlier than females’. Our results are consistent with other recent studies in darters and suggest that the coevolution of female preferences and male ornaments may not best explain the earliest stages of behavioral isolation in this lineage.     

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.     

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.     

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.     

Sexual differences in certain forms of motor asymmetry in rats

Using three tests, studies have been made on locomotor lateralization in Wistar rats. It was shown that rat prefer the left paw for obtaining food from a tube, this preference being more pronounced in females. Females are more asymmetric also in preference of a side turn in a labyrinth: they choose the right side. In males, the preference of the right and left turns, as well as the absence of locomotor asymmetry were almost equal. These data are rather similar to those obtained for human subjects. Studies on the direction of rotation of rats after apomorphine injections did not reveal any sex differences or populational preferences.     

Female swamp sparrows do not show evidence of discriminating between the songs of peak‐aged and senescent males

Sexual selection theory predicts that females face contrasting selection pressures when choosing the age of their mate. On the one hand, older males have demonstrated their ability to survive and they may be more experienced than younger males. At the same time, however, younger males are expected to have accumulated fewer deleterious mutations in their germline as compared to older males. These contrasting pressures on female preference may result in a preference for intermediate‐aged males. A preference for males of a particular age can only be expressed, however, if females are able to identify males of different ages. We have previously shown that male swamp sparrows display age‐related changes in vocal quality, such that males display sharp increases in vocal quality in early adulthood, followed by gradual senescent declines thereafter. We have also shown that territorial males discriminate these within‐individual differences, giving stronger aggressive responses to songs of peak‐aged males than to those of senescent males. Here, we use a copulation solicitation assay to test whether females also discriminate these within‐signaler markers of senescence in song. Contrary to our prediction, females did not show any evidence of discriminating between songs recorded from peak‐aged males as compared to songs from the same males following song senescence. We suggest that this difference in demonstrated discrimination between males and females may be the result of the two sexes attending to different song characteristics.     

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.     

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.