The evolution of self-control.

Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.

Delay of gratification by orangutans (Pongo pygmaeus) in the accumulation task.

There is considerable evidence indicating that chimpanzees can delay gratification for extended time intervals, particularly in the accumulation task in which food items accumulate within a participant's reach until the participant begins to consume them. However, there is limited evidence that other ape species might also exhibit this capacity, despite there being a number of similar studies indicating that nonape species (e.g., monkeys and birds) can delay gratification, but not for nearly as long as chimpanzees. To help define the taxonomic distribution of delay of gratification behavior in the order Primates, we tested 6 orangutans in the current experiments and compared their performance with comparable data from a previous study with capuchin monkeys. We varied delay length and visibility of the items that were still available for accumulation to determine the impact of these factors on performance. Species differences on the accumulation task emerged when comparing the current data to data from a previous study. Orangutans outperformed capuchin monkeys, suggesting that ape species may generally show better delay of gratification and delay maintenance abilities than monkeys. However, more studies are necessary to rule out alternative hypotheses on the distribution of delay maintenance abilities across primate species.

Gorillas' use of the escape response in object choice memory tests.

The ability to monitor and control one's own cognitive states, metacognition, is crucial for effective learning and problem solving. Although the literature on animal metacognition has grown considerably during last 15 years, there have been few studies examining whether great apes share such introspective abilities with humans. Here, we tested whether four gorillas could meet two criteria of animal metacognition, the increase in escape responses as a function of task difficulty and the chosen-forced performance advantage. During testing, the subjects participated in a series of object choice memory tests in which a preferable reward (two grapes) was placed under one of two or three blue cups. The apes were required to correctly select the baited blue cup in this primary test. Importantly, the subjects also had an escape response (a yellow cup), where they could obtain a secure but smaller reward (one grape) without taking the memory test. Although the gorillas received a relatively small number of trials and thus experienced little training, three gorillas significantly declined the memory tests more often in difficult trials (e.g., when the location of the preferred reward conflicted with side bias) than in easy trials (e.g., when there was no such conflict). Moreover, even when objective cues were eliminated that corresponded to task difficulty, one of the successful gorillas showed evidence suggestive of improved memory performance with the help of escape response by selectively avoiding trials in which he would be likely to err before the memory test actually proceeded. Together, these findings demonstrate that at least some gorillas may be able to make optimal choices on the basis of their own memory trace strength about the location of the preferred reward.

The effect of body orientation on judgments of human visual attention in western lowland gorillas (Gorilla gorilla gorilla).

The ability to appreciate the attentional states of others is an important element used in discerning the presence of Theory of Mind in an individual. Whether primates are able to recognize attention, and further, use such information to predict behavior, remains contentious. In this study, six western lowland gorillas (Gorilla gorilla gorilla) were tested under different conditions that aimed to investigate not only their understanding of what a human experimenter sees, but also what information they use to make these judgments. In all experiments the gorillas selected between two human experimenters, one who could see them and one who could not. In Experiment 1 the gorillas' performance was significantly above chance in conditions where they judged body orientation, head orientation, and eyes. Experiment 2 showed that the gorillas' initial judgments of attention may be based on body orientation rather than facial cues. Experiment 3 demonstrated that the gorillas were better able to utilize facial cues in some conditions, when these cues were paired with a more neutral body orientation. These findings suggest that the gorillas are using a hierarchical approach to determining visual attention, by making an immediate decision based on the most salient cue-body orientation. However, when body orientation is more ambiguous their ability to judge the less prominent cues of the face and eyes does improve.

Constructive and deconstructive tool modification by chimpanzees (Pan troglodytes).

Nine chimpanzees (Pan troglodytes) were tested for their ability to assemble or disassemble the appropriate tool to obtain a food reward from two different apparatus. In its deconstructed form, the tool functioned as a probe for one apparatus. In its constructed form, the tool functioned as a hook, appropriate for a second apparatus. Each subject completed four test trials with each apparatus type. Tool types were randomized and counter-balanced between the two forms. Results demonstrated that adult and juvenile chimpanzees (N = 7) were successful with both tool types, while two infant chimpanzees performed near chance. Off-line video analyses revealed that tool modifications followed by attempted solutions by the adults and juveniles were typically correct on the first attempt. Neither infant was successful in modifying tools correctly on the first attempt over all eight trials. The older chimpanzees' ability to modify the appropriate tool consistently prior to use indicates an immediate recognition of the functional attributes necessary for the successful use of tool types on each apparatus, and represents a non-replication of a previously reported study by Povinelli.