William D. Hopkins

Asymmetric Broca's area in great apes

Brodmanns area 44 delineates part of Brocas area within the inferior frontal gyrus of the human brain and is a critical region for speech production, being larger in the left hemisphere than in the right — an asymmetry that has been correlated with language dominance. Here we show that there is a similar asymmetry in this area, also with left-hemisphere dominance, in three great ape species (Pan troglodytes, Pan paniscus and Gorilla gorilla). Our findings suggest that the neuroanatomical substrates for left-hemisphere dominance in speech production were evident at least five million years ago and are not unique to hominid evolution.

Hand preferences for a coordinated bimanual task in 110 chimpanzees (Pan troglodytes): cross-sectional analysis.

Hand preference for a coordinated bimanual task was assessed in a sample of 110 chimpanzees (Pan troglodytes). Subjects were presented with opaque tubes, the inside of which was coated with peanut butter. The hand and finger used to extract the peanut butter was recorded in 2 test sessions. A population right-hand bias was found. Juvenile and adolescent subjects were less lateralized than adults. All subjects primarily used their index finger to extract peanut butter. Use of the index finger was higher in adults and adolescents compared with juveniles. The results suggest that coordinated bimanual tasks (a) elicit strong hand preferences at the individual level and (b) elicit population level right-handedness.

Understanding the Point of Chimpanzee Pointing: Epigenesis and Ecological Validity

Pointing has long been considered to be a uniquely human, universal, and biologically based gesture. However, pointing emerges spontaneously, without explicit training, in captive chimpanzees. Because pointing is commonplace in captive chimpanzees and virtually absent in wild chimpanzees, and because both captive and wild chimpanzees are sampled from the same gene pool, pointing by captive apes is attributable to environmental influences on communicative development. If pointing by captive chimpanzees is so variably expressed in different rearing environments, this suggests that pointing by humans may also be attributable to situational factors that make pointing effective in certain developmental contexts.

Comparative and familial analysis of handedness in great apes.

Historically, population-level handedness has been considered a hallmark of human evolution. Whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. This paper summarizes published data on handedness in great apes. Comparative analysis indicated that chimpanzees and bonobos show population-level right handedness, whereas gorillas and orangutans do not. All ape species showed evidence of population-level handedness when considering specific tasks. Familial analyses in chimpanzees indicated that offspring and maternal (but not paternal) handedness was significantly positively correlated, but this finding was contingent upon the classification criteria used to evaluate hand preference. Overall, the proportion of right handedness is lower in great apes compared with humans, and various methodological and theoretical explanations for this discrepancy are discussed.

Referential communication by chimpanzees (Pan troglodytes)

Two experiments were conducted to assess the referential function of chimpanzee (Pan troglodytes) gestures to obtain food. The chimpanzees received 1 trial per condition. In Experiment 1 (N = 101), in full view of the chimpanzee, a banana was placed on top of 1 of 2 inverted buckets or was hidden underneath 1 of the buckets. In Experiment 2 (N = 35), 4 conditions were presented in constant order: (a) no food, no observer; (b) no food, observer present; (c) food present, no observer; and (d) food present, observer present. Gestures and visual orienting were used socially and referentially. The capacity for nonverbal reference may predate the Hominidae-Pongidae split, and the development of nonverbal reference may be independent of human species-specific adaptations for speech.

Hand use and gestural communication in chimpanzees (Pan troglodytes).

Hand use in gestural communication was examined in 115 captive chimpanzees (Pan troglodytes). Hand use was measured in subjects while they gestured to food placed out of their reach. The distribution of hand use was examined in relation to sex, age, rearing history, gesture type, and whether the subjects vocalized while gesturing. Overall, significantly more chimpanzees, especially females and adults, gestured with their right than with their left hand. Foods begs were more lateralized to the right hand than pointing, and a greater prevalence of right-hand gesturing was found in subjects who simultaneously vocalized than those who did not. Taken together, these data suggest that referential, intentional communicative behaviors, in the form of gestures, are lateralized to the left hemisphere in chimpanzees.

On the Other Hand: Statistical Issues in the Assessment and Interpretation of Hand Preference Data in Nonhuman Primates

I describe methodological and statistical issues in the assessment of hand preference in nonhuman primates and discuss them in the context of a recent paper by McGrew and Marchant (1997) in which they conclude that there is no convincing evidence of population-level hand preferences in nonhuman primates. The criteria used by them to evaluate individual and population-level hand preferences are flawed, which results in an oversimplification of findings in nonhuman primates. I further argue that the classification schema used by McGrew and Marchant (1997) to compare hand preference distributions between species is theoretically weak and does not offer a meaningful way to compare human and nonhuman primate handedness.

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.

Individual and setting differences in the hand preferences of chimpanzees (Pan troglodytes): A critical analysis and some alternative explanations

Several recent papers have been critical at a theoretical and empirical level of the evidence of population-level right-handedness in chimpanzees and other great apes. For example, Palmer (2002) has recently argued that the evidence of population-level handedness in chimpanzees is weak because there are sampling biases in the data. McGrew and Marchant (1997) argue that all the evidence of right-handedness in apes is from captive animals and therefore the observed phenomenon has little ecological validity. In this paper, we address recent issues regarding the presentation and interpretation of other hand preference data and argue that chimpanzees are right-handed for some measures. We further argue that purported differences in hand use between wild and captive chimpanzees due to rearing environments are unfounded and we emphasise that more cooperative work between researchers working in captive and feral populations is needed to facilitate collection of data on common measures of hand preference.

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.