Chimpanzees (Pan troglodytes) Are Predominantly Right-Handed: Replication in Three Populations of Apes.

Population-level right-handedness has historically been considered a hallmark of human evolution. Even though recent studies in chimpanzees (Pan troglodytes) have demonstrated population-level right-handedness for certain behaviors, some have questioned the validity and consistency of these findings by arguing that reported laterality effects are specific to certain colonies of apes and to those chimpanzees reared by humans. The authors report evidence of population-level right-handedness in 3 separate colonies of chimpanzees. Moreover, handedness in the 3 colonies was unrelated to the proportion of subjects that were raised by humans. This is the strongest evidence to date that population-level handedness is evident in chimpanzees and is not an artifact of human rearing. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Handedness for tool use correlates with cerebellar asymmetries in chimpanzees (Pan troglodytes).

Recent studies have shown that great ape species possess patterns of macrostructural neocortical asymmetries that are similar to those found in humans. However, little is known about the asymmetry of subcortical structures in great apes. To address this lack of data, the authors assessed left-right asymmetry of the anterior and posterior aspects of cerebellum from MRI brain scans of 53 chimpanzees (Pan troglodytes). No population-level bias was found for either the anterior or the posterior region of the cerebellum. However, a significant inverse association was found in the asymmetry quotients of the anterior and posterior regions, indicating that the cerebellum was torqued at the individual level. Additionally, handedness for tool use but not other measures was associated with variation in cerebellar asymmetries. Last, older chimpanzees had a smaller cerebellum after brain volume was adjusted for. The results are discussed in the context of brain changes in primate evolution related to tool use. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A Meta-Analysis of Primate Hand Preferences, Particularly for Reaching.

P. F. MacNeilage, M. G. Studdert-Kennedy, and B. Lindblom (1987) proposed a progression for handedness in primates that was supposed to account for the evolution of a right bias in human handedness. To test this proposal, the authors performed meta-analyses on 62 studies that provided individual data (representing 31 species: 9 prosimians, 6 New World monkeys, 10 Old World monkeys, 2 lesser apes, and 4 greater apes), of the 118 studies of primate handedness published since 1987. Although evidence of a population-level left-handed bias for prosimians and Old World monkeys supports P. F. MacNeilage et al., the data from apes, New World monkeys, and individual species of prosimians and New World monkeys do not. Something other than primate handedness may have been the evolutionary precursor of the right bias in hand-use distribution among hominids. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Assessment of shoulder dimensions and angles of porcelain bonded to metal crown preparations

This research examined hand preference for a bimanual task in 45 tufted capuchin (Cebus apella) and 55 rhesus macaque (Macaca mulatta) monkeys. Investigators presented subjects with plastic tubes lined with food and noted which hand the animals used to hold the tubes and which hand the animals used to remove the food. Several significant findings emerged from this investigation. First, rhesus macaques, but not tufted capuchins, exhibited a population-level bias toward use of the right hand (although the difference in direction of hand preference between species was not significant). Second, capuchins exhibited greater hand preference strength than did macaques. Third, among capuchins, but not among macaques, hand preference strength was greater for adults than for immatures. Finally, both species used their index digit to remove food most frequently when compared with other digits. Findings of hand preference direction and strength in this study were compared with other findings noted for chimpanzees which performed a bimanual tube task in a previous study. The authors conclude that using the same procedure to compare hand preference across species represents a powerful research tool that can lead to a more complete understanding of the evolution and ontogenesis of primate handedness.     

Heritability of hand preference in chimpanzees (Pan troglodytes): Evidence from a partial interspecies cross-fostering study.

Heritability of hand preference was tested in a sample of 188 chimpanzees (Pan troglodytes). Hand preference was measured by coordinated bimanual actions, and concordance percentages were compared between parents and offspring and siblings. Among siblings, concordance percentages were compared for dyads in which both individuals were raised by chimpanzees, both were raised by humans, or 1 was raised in each environment. The results indicated population-level right hand preferences for coordinated bimanual actions. There were no significant associations in hand preference between parents and offspring. In full and maternal half siblings, concordance in hand preference was significantly greater than chance in mother and human-reared individuals but not in cross-fostered dyads. The cumulative results suggest that the direction of hand preference is heritable in chimpanzees but the mechanism of transmission is not genetic. Several environmental explanations are proposed to explain the findings, including the potential role of maternal cradling bias and in utero fetal position. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preference for a bimanual task in tufted capuchins (Cebus apella) and rhesus macaques (Macaca mulatta).

Examined hand preference for a bimanual task in 45 tufted capuchin (Cebus apella) and 55 rhesus macaque (Macaca mulatta) monkeys. Investigators presented subjects with plastic tubes lined with food and noted which hand the animals used to hold the tubes and which hand the animals used to remove the food. Several significant findings emerged from this investigation. First, rhesus macaques, but not tufted capuchins, exhibited a population-level bias toward use of the right hand (although the difference in direction of hand preference between species was not significant). Second, capuchins exhibited greater hand preference strength than did macaques. Third, among capuchins, but not among macaques, hand preference strength was greater for adults than for immatures. Finally, both species used their index digit to remove food most frequently when compared with other digits. Findings of hand preference direction and strength in this study were compared with other findings noted for chimpanzees which performed a bimanual tube task in a previous study. The authors conclude that using the same procedure to compare hand preference across species represents a powerful research tool that can lead to a more complete understanding of the evolution and ontogenesis of primate handedness. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand preference for food processing in wild western lowland gorillas (gorilla gorilla gorilla).

Western lowland gorillas (Gorilla gorilla gorilla) have not previously been represented in studies of laterality in wild great apes. The discovery of swampy clearings frequented by gorillas in northern Congo has provided the first opportunity to redress this imbalance. Hand preference data are presented from 33 gorillas in seated and standing postures, covering the procurement and processing of 2 to 4 plant species. Levels of hand preference exhibited were low. When data from all postures and plant species were pooled, 33% of gorillas showed hand preferences in excess of chance. In the standing posture, more gorillas exhibited significant left-hand preferences than right, but an overall population-level bias was not evident. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Asymmetries in the Hippocampus and Amygdala of Chimpanzees (Pan troglodytes).

Magnetic resonance imaging was used to measure the hippocampal and amygdalar volumes of 60 chimpanzees (Pan troglodytes). An asymmetry quotient (AQ) was then used to calculate the asymmetry for each of the structures. A one-sample t test indicated that there was a population-level right hemisphere asymmetry for the hippocampus. There was no significant population-level asymmetry for the amygdala. An analysis of variance using sex and rearing history as between-group variables showed no significant main effects or interaction effects on the AQ scores; however, males were more strongly lateralized than females. Several of these findings are consistent with results found in the human literature. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Posture and reaching in chimpanzees (Pan troglodytes) and orangutans (Pongo pygmaeus).

Hand preference in reaching was assessed under 2 postural adjustment conditions in 40 chimpanzees (Pan troglodytes) and 6 orangutans (Pongo pygmaeus). The postural conditions were quadrupedal and upright, during which reaching for food was scored on a minimum of 50 trials. Results indicated no population preference during quadrupedal reaching, but a right-hand population preference was found during upright reaching. There were no significant effects of age or sex on either the direction or strength of hand preference. Early rearing history affected the strength of hand preference but not direction. The results suggest that posture is an important factor in the assessment of hand preference in great apes and may have important evolutionary consequences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The use of bouts and frequencies in the evaluation of hand preferences for a coordinated bimanual task in chimpanzees (Pan troglodytes): An empirical study comparing two different indices of laterality.

Hand preferences for a coordinated bimanual task were assessed in 109 chimpanzees (Pan troglodytes). Hand preference was evaluated for 4 test sessions using bouts and frequencies of hand use to compare the sensitivity of each level of analysis in evaluating individual variation in handedness. Overall, significant population-level right-handedness was found using several different measures of hand use. Handedness indices based on bouts and frequencies were highly and significantly correlated. Moreover, hand preferences were consistent across tests despite efforts to situationally bias preference during each test. Taken together, these data do not support the view that bouts are a better level of analysis for evaluating hand preference. The results further suggest that hand preferences for coordinated bimanual actions are not influenced by situational factors and may reflect an inherent specialization of the left hemisphere for motor skill. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Postural effects on manual reaching laterality in squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus).

Squirrel monkeys (Saimiri sciureus) and cotton-top tamarins (Saguinus oedipus) made unimanual food reaches from either a horizontal, quadrupedal posture or a vertical, clinging posture. No population-level handedness occurred in either species. However, in both species, directional lateral preferences weakly expressed for reaching from the stable quadrupedal posture were intensified in the vertical cling posture. This phenomenon, which is designated as soft handedness, may have been an evolutionary precursor to population-level handedness. Right or left turning by the squirrel monkeys before reaching closely predicted use of the right or left hand. However, the magnitude of the association decreased in the more highly lateralized vertical cling condition. This result suggests that as lateral hand preference increases, hand use may become increasingly independent of constraints from prior behavioral and environmental influences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Developmental issues in the evolution of the mind.

D. J. Povinelli (see record 1993-35769-001) did not integrate theory-of-mind (TOM) literature with that in other cognitive domains, and did not reference other investigators' research on TOM in great apes. Povinelli's claim that most chimpanzees do not display evidence of self-recognition until age 6–8 yrs is refuted with reference to this research. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Token transfers among great apes (Gorilla gorilla, Pongo pygmaeus, Pan paniscus, and Pan troglodytes): Species differences, gestural requests, and reciprocal exchange.

Great apes appear to be the nonhuman primates most capable of recognizing trading opportunities and engaging in transfers of commodities with conspecifics. Spontaneous exchange of goods between them has not yet been reported. We tested gorillas (Gorilla gorilla), orangutans (Pongo pygmaeus), bonobos (Pan paniscus), and chimpanzees (Pan troglodytes) in a token-exchange task involving two conspecifics and a human experimenter. Tested in pairs, subjects had to exchange tokens with a partner to obtain food from the experimenter. We observed 4, 5, 264, and 328 transfers of tokens in gorillas, chimpanzees, orangutans, and bonobos, respectively. Most gifts were indirect in gorillas, chimpanzees, and bonobos, whereas most were direct in orangutans. The analysis showed no evidence of calculated reciprocity in interactions. A main finding of the study was the high rate of repeated gifts and begging gestures recorded in orangutans. This raises the question of the meaning of pointing in great apes and their ability to understand the communicative intent of others. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Physical intuitions about support relations in monkeys (Macaca fuscata) and apes (Pan troglodytes).

Nonhuman primates, like humans, have demonstrated various physical intuitions. Cacchione and Krist (2004) examined chimpanzees' intuitions about support relations with the violation-of-expectation task. They reported that the chimpanzees possessed intuitions about support, but their intuitions differed from those of humans in part; they were sensitive to “contact/no-contact” and “amount of contact” but not “type of contact” rule. To further explore intuitions about support in nonhuman primates, we conducted similar experiments on monkeys (Japanese monkeys) and apes (chimpanzees). In three experiments, we presented physically possible and impossible events of different support relations to the participants and measured their looking times. The results reveal that the chimpanzees and monkeys detect the violations of “contact/no-contact” and “amount of contact” but not “type of contact” variable. Therefore, the apes and monkeys possess similar intuitions; however, these intuitions differ in part from those of humans. The present study provides new and corroborative evidence of intuitions about support in nonhuman primates. However, this again leads to the question of distinctive understanding about support relations among primate species. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

The magic cup: Great apes and domestic dogs (Canis familiaris) individuate objects according to their properties.

Despite current interest in dog (Canis familiaris) cognition, very little is known about how dogs represent objects and how they compare with other species, such as the great apes. Therefore, we investigated how dogs and great apes (chimpanzees [Pan troglodytes], bonobos [Pan paniscus], orangutans [Pongo pygmaeus], gorillas [Gorilla gorilla]) individuate objects in a classical violation of expectation paradigm. We used a container (magic cup) with a double bottom that allowed us to change the type of food that subjects had seen being placed in the container. Using a 2 × 2 design, we varied whether subjects received a generally preferred food and whether the food was substituted (surprise trials) or not (baseline trials). Apes showed increased begging and looking behaviors and dogs showed increased smelling behavior. Both species stayed near the experimenter more frequently in the surprise trials compared with baseline trials. Both species reacted to positive (i.e., good food substituted for bad food) and negative (i.e., bad food substituted for good food) surprises. These results suggest that apes and dogs were able to individuate objects according to their properties or type in comparable ways. In addition, we looked for frustration and elation effects, but subjects' behaviors were not influenced by the food they saw and which they received in previous trials. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Aping expressions? Chimpanzees produce distinct laugh types when responding to laughter of others.

Humans have the ability to replicate the emotional expressions of others even when they undergo different emotions. Such distinct responses of expressions, especially positive expressions, play a central role in everyday social communication of humans and may give the responding individuals important advantages in cooperation and communication. The present work examined laughter in chimpanzees to test whether nonhuman primates also use their expressions in such distinct ways. The approach was first to examine the form and occurrence of laugh replications (laughter after the laughter of others) and spontaneous laughter of chimpanzees during social play and then to test whether their laugh replications represented laugh-elicited laugh responses (laughter triggered by the laughter of others) by using a quantitative method designed to measure responses in natural social settings. The results of this study indicated that chimpanzees produce laugh-elicited laughter that is distinct in form and occurrence from their spontaneous laughter. These findings provide the first empirical evidence that nonhuman primates have the ability to replicate the expressions of others by producing expressions that differ in their underlying emotions and social implications. The data further showed that the laugh-elicited laugh responses of the subjects were closely linked to play maintenance, suggesting that chimpanzees might gain important cooperative and communicative advantages by responding with laughter to the laughter of their social partners. Notably, some chimpanzee groups of this study responded more with laughter than others, an outcome that provides empirical support of a socialization of expressions in great apes similar to that of humans. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Chimpanzee (Pan troglodytes) handedness: Variability across multiple measures of hand use.

This study examined intertask consistency in handedness across multiple measures of hand use in a sample of 187 chimpanzees (Pan troglodytes). Hand preferences for 2 to 6 measures were collected from the sample, and hand preference scores were derived on the basis of the individual hand preferences for each measure. Seven of 15 possible intratask correlations were significant, with some degree of clustering depending on the motor demands of the tasks. Two overall measures of handedness revealed population-level right-handedness in the chimpanzees, although the degree of bias was reduced for chimpanzees tested on more than 3 measures of hand use. The results are interpreted in the context of several recent studies that proposed theoretical models of handedness in nonhuman primates. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Note on hand use in the manipulation of joysticks by rhesus monkeys (Macaca mulatta) and chimpanzees (Pan troglodytes).

In a recent article MacNeilage, Studdert-Kennedy, and Lindblom (1987) proposed that nonhuman primate handedness may be contingent on the specific task requirements with visual–spatial tasks yielding left-hand preferences and fine motor tasks producing right-hand preferences. This study reports hand preferences in the manipulation of joysticks by 2 rhesus monkeys (Macaca mulatta) and 3 chimpanzees (Pan troglodytes). Reach data were collected on these same subjects and served as a basis for comparison with preference data for manipulation of the joystick. The data indicated that all 5 subjects demonstrated significant right-hand preferences in manipulating the joystick. In contrast, no significant hand preferences were found for the reach data. Reaction time (RT) data also indicated that the right could perform a perceptual–motor task better than the left hand in all 5 subjects. Overall, the data indicate that reach tasks may not be sensitive enough measures to produce reliable hand preferences, whereas tasks that assess fine motor control produce significant hand preferences. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Methodology of motor skill and laterality: new test of hand preference in Macaca nemestrina

Ten female pig-tailed macaques (Macaca nemestrina) were tested for hand preference and hand skill (i.e., speed of performance and error rate). The experimental task was naturalistic, calling for adaptive manual skills and fine manipulation: the monkeys had to remove small food rewards embedded in a vertical array, and precision opposition of thumb and forefinger was needed to extract each pellet. Each monkey was tested 10 times on 10 different days. The results indicated individual hand preference rather than population-level handedness; however, a tendency toward right predominance was found. The results on hand skill showed a relation between error rate and hand preference, as the preferred hand made fewer errors. A different and unexpected finding was obtained when skill was evaluated in terms of speed of performance: in adult subjects the left hand was quicker than the right. Therefore, different kinds of skill showed different patterns in relation to hand preference.