Hani D. Freeman

Personality in nonhuman primates: a review and evaluation of past research.

Scientific reports of personality in nonhuman primates are now appearing with increasing frequency across a wide range of disciplines, including psychology, anthropology, endocrinology, and zoo management. To identify general patterns of research and summarize the major findings to date, we present a comprehensive review of the literature, allowing us to pinpoint the major gaps in knowledge and determine what research challenges lay ahead. An exhaustive search of five scientific databases identified 210 relevant research reports. These articles began to appear in the 1930s, but it was not until the 1980s that research on primate personality began to gather pace, with more than 100 articles published in the last decade. Our analyses of the literature indicate that some domains (e.g., sex, age, rearing conditions) are more evenly represented in the literature than are others (e.g., species, research location). Studies examining personality structure (e.g., with factor analysis) have identified personality dimensions that can be divided into 14 broad categories, with Sociability, Confidence/Aggression, and Fearfulness receiving the most research attention. Analyses of the findings pertaining to inter‐rater agreement, internal consistency, test–retest reliability, generally support not only the reliability of primate personality ratings scales but also point to the need for more psychometric studies and greater consistency in how the analyses are reported. When measured at the level of broad dimensions, Extraversion and Dominance generally demonstrated the highest levels of inter‐rater reliability, with weaker findings for the dimensions of Agreeableness, Emotionality, and Conscientiousness. Few studies provided data with regard to convergent and discriminant validity; Excitability and Dominance demonstrated the strongest validity coefficients when validated against relevant behavioral criterion measures. Overall, the validity data present a somewhat mixed picture, suggesting that high levels of validity are attainable, but by no means guaranteed. Discussion focuses on delineating major theoretical and empirical questions facing research and practice in primate personality. Am. J. Primatol. 72:653–671, 2010.

The Distribution and Development of Handedness for Manual Gestures in Captive Chimpanzees (Pan troglodytes)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, noncommunicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.

Now you see me, now you don't: evidence that chimpanzees understand the role of the eyes in attention

Chimpanzees appear to understand something about the attentional states of others; in the present experiment, we investigated whether they understand that the attentional state of a human is based on eye gaze. In all, 116 adult chimpanzees were offered food by an experimenter who engaged in one of the four experimental manipulations: eyes closed, eyes open, hand over eyes, and hand over mouth. The communicative behavior of the chimpanzees was observed. More visible behaviors were produced when the experimenters eyes were visible than when the experimenters eyes were not visible. More vocalizations were produced when the experimenters eyes were closed than when they were open, but there were no differences in other attention getting behaviors. There was no effect of age or rearing history. The results suggest that chimpanzees use the presence of the eyes as a cue that their visual gestures will be effective.

Developing a Comprehensive and Comparative Questionnaire for Measuring Personality in Chimpanzees Using a Simultaneous Top-Down/Bottom-Up Design

One effective method for measuring personality in primates is to use personality trait ratings to distill the experience of people familiar with the individual animals. Previous rating instruments were created using either top‐down or bottom‐up approaches. Top‐down approaches, which essentially adapt instruments originally designed for use with another species, can unfortunately lead to the inclusion of traits irrelevant to chimpanzees or fail to include all relevant aspects of chimpanzee personality. Conversely, because bottom‐up approaches derive traits specifically for chimpanzees, their unique items may impede comparisons with findings in other studies and other species. To address the limitations of each approach, we developed a new personality rating scale using a combined top‐down/bottom‐up design. Seventeen raters rated 99 chimpanzees on the new 41‐item scale, with all but one item being rated reliably. Principal components analysis, using both varimax and direct oblimin rotations, identified six broad factors. Strong evidence was found for five of the factors (Reactivity/Undependability, Dominance, Openness, Extraversion, and Agreeableness). A sixth factor (Methodical) was offered provisionally until more data are collected. We validated the factors against behavioral data collected independently on the chimpanzees. The five factors demonstrated good evidence for convergent and predictive validity, thereby underscoring the robustness of the factors. Our combined top‐down/bottom‐up approach provides the most extensive data to date to support the universal existence of these five personality factors in chimpanzees. This framework, which facilitates cross‐species comparisons, can also play a vital role in understanding the evolution of personality and can assist with husbandry and welfare efforts. Am. J. Primatol. 75:1042–1053, 2013.

Factors influencing the prevalence and handedness for throwing in captive chimpanzees (Pan troglodytes).

Humans throw right-handed, and it has been suggested that the neurophysiological demands of aimed throwing may have served as a precursor to the evolution of left hemisphere specialization for linguistic functions. Although there are descriptions of throwing by wild and captive chimpanzees (Pan troglodytes), systematic observations of aimed throwing and handedness have not been reported. In this article, evidence of population-level right-handedness for throwing is reported in 2 samples of captive chimpanzees. It is further reported that right-handed throwing is more pronounced than other measures of handedness in captive chimpanzees. The implications of these findings are discussed in the context of theories relating throwing to the evolution of lateralization for language functions.

Squirrel monkeys' response to inequitable outcomes indicates a behavioural convergence within the primates

Although several primates respond negatively to inequity, it is unknown whether this results from homology or convergent processes. Behaviours shared within a taxonomic group are often assumed to be homologous, yet this distinction is important for a better understanding of the function of the behaviour. Previous hypotheses have linked cooperation and inequity responses. Supporting this, all species in which inequity responses have been documented are cooperative. In this study, we tested this hypothesis by investigating the response to inequity in squirrel monkeys, which share a phylogenetic family with capuchin monkeys, but do not cooperate extensively. Subjects exchanged tokens to receive food rewards in conditions in which the level of effort required and reward received varied. Squirrel monkeys did not respond negatively to inequity. However, the monkeys were sensitive to the variation present in the task; male subjects showed a contrast effect and, as in previous studies, subjects were more sensitive to differences in reward in the context of a task than when rewards were given for free. Taken with other results, these results support the hypothesis that a negative response to inequity evolved convergently in primates, probably as a mechanism for evaluating outcomes relative to ones partners in cooperative species.

Chimpanzees are right‐handed when recording bouts of hand use

Whether nonhuman primates exhibit population‐level handedness remains a topic of considerable debate. Previous research has shown that chimpanzees are right‐handed when frequencies of hand use are recorded but some have questioned the validity of this approach. In this study, we evaluated handedness in 180 captive chimpanzees for a task measuring bimanual actions. Bouts rather than frequency of hand use were recorded in each subject. Population‐level right‐handedness was found using both continuous and nominal scales of measurement. Neither sex nor rearing history had a significant effect on hand use. These results indicate that chimpanzees are right‐handed, even when using a more conservative measure of handedness. Limitations in the use of bouts in handedness assessment are also discussed.

Left Hemisphere Specialization for Oro-Facial Movements of Learned Vocal Signals by Captive Chimpanzees

Background The left hemisphere of the human brain is dominant in the production of speech and signed language. Whether similar lateralization of function for communicative signal production is present in other primates remains a topic of considerable debate. In the current study, we examined whether oro-facial movements associated with the production of learned attention-getting sounds are differentially lateralized compared to facial expressions associated with the production of species-typical emotional vocalizations in chimpanzees. Methodology/ Principal Findings Still images captured from digital video were used to quantify oro-facial asymmetries in the production of two attention-getting sounds and two species-typical vocalizations in a sample of captive chimpanzees. Comparisons of mouth asymmetries during production of these sounds revealed significant rightward biased asymmetries for the attention-getting sounds and significant leftward biased asymmetries for the species-typical sounds. Conclusions/Significance These results suggest that the motor control of oro-facial movements associated with the production of learned sounds is lateralized to the left hemisphere in chimpanzees. Furthermore, the findings suggest that the antecedents for lateralization of human speech may have been present in the common ancestor of chimpanzees and humans ∼5 mya and are not unique to the human lineage.

Handedness and Grooming in Pan troglodytes: Comparative Analysis Between Findings in Captive and Wild Individuals

Grooming is a complex set of motor actions, common in highly social primates. We tested for asymmetries in hand use during unimanual and bimanual allogrooming in 215 captive chimpanzees. In addition to hand use, we coded in the ethogram whether the manual grooming action co-occurred with the use of the mouth. Overall, grooming did not elicit strong handedness at the individual level, but there is a small yet significant population-level right-hand bias for bimanual grooming. Mouth use during grooming had no influence on hand use. A comparison of the findings with previously published data on handedness for grooming in wild chimpanzees suggests that wild apes are more right-handed than captive individuals are for allogrooming. Collectively, the results suggest that role differentiation of the hands is an important factor in the assessment of handedness for grooming, and perhaps additional manual actions of chimpanzees and other primates.

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.