Mirror self-recognition and its relationship to social cognition in chimpanzees.

Chimpanzees and humans are capable of recognizing their own reflection in mirrors. Little is understood about the selective pressures that led to this evolved trait and about the mechanisms that underlie it. Here, we investigated the hypothesis that mirror self-recognition in chimpanzees is the byproduct of a developed form of self-awareness that was naturally selected for its adaptive use in social cognitive behaviors. We present here the first direct attempt to assess the social cognition hypothesis by analyzing the association between mirror self-recognition in chimpanzees, as measured by a mirror-mark test, and their performance on a variety of social cognition tests. Consistent with the social cognition hypothesis, chimpanzees who showed evidence of mirror self-recognition in the mark test tended to perform significantly better on the social cognition tasks than those who failed the mark test. Additionally, the data as a whole fit the social cognition hypothesis better than the main competing hypothesis of mirror self-recognition in great apes, the secondary representation hypothesis. Our findings strongly suggest that the evolutionary origins of great apes' and humans' capacity to understand ourselves, as revealed by our capacity to recognize ourselves in mirrors, are intimately linked to our ability to understand others.

Chimpanzees gesture to humans in mirrors: using reflection to dissociate seeing from line of gaze.

There is much experimental evidence suggesting that chimpanzees understand that others see. However, previous research has never experimentally ruled out the alternative explanation that chimpanzees are just responding to the geometric cue of 'direct line of gaze', the observable correlate of seeing in others. Here, we sought to resolve this ambiguity by dissociating seeing from direct line of gaze using a mirror. We investigated the frequency of chimpanzees' visual gestures towards a human experimenter who could see them (as a result of looking into a mirror) but who lacked a direct line of gaze to them (as a result of having his/her head turned away). Chimpanzees produced significantly more visual gestures when the experimenter could see them than when he/she could not, even when the experimenter did not have a direct line of gaze to them. Results suggest that chimpanzees, through a possible process of experience projection based on their own prior experience with mirrors, infer that an experimenter looking at the mirror can see them. We discuss our results in relation to the theory of mind hypothesis that chimpanzees understand seeing in others, and we evaluate possible alternative low-level explanations.

I know you see it wrong! Children use others' false perceptions to predict their behaviors.

Research on children's ability to attribute false mental states to others has focused exclusively on false beliefs. We developed a novel paradigm that focuses instead on another type of false mental state: false perceptions. From approximately 4years of age, children begin to recognize that their perception of an illusory object can be at odds with its true properties. Our question was whether they also recognize that another individual viewing the object will similarly experience a false perception. We tested 33 preschool children with a task in which distorting lenses caused a small object to appear large and a large object to appear small. To succeed, children needed to recognize that a naive agent would falsely perceive the relative size of the objects and to correctly anticipate the agent's actions on that basis. Children performed significantly better than chance in our false perception test, and there was a developmental progression in performance from 4 to 5years of age similar to that seen in standard false belief tests. Our findings demonstrate that preschool children are capable of understanding that other individuals will be perceptually misled by illusory objects and that these false perceptions will influence their actions in predictable ways.

Smoke and mirrors: Testing the scope of chimpanzees' appearance-reality understanding.

The ability to make appearance-reality (AR) discriminations is an important higher-order cognitive adaptation in humans but is still poorly understood in our closest primate relatives. Previous research showed that chimpanzees are capable of AR discrimination when choosing between food items that appear, due to the effects of distorting lenses, to be smaller or larger than they actually are (Krachun, Call, & Tomasello, 2009). In the current study, we investigated the scope and flexibility of chimpanzees' AR discrimination abilities by presenting them with a wider range of illusory stimuli. In addition to using lenses to change the apparent size of food items (Experiment 1), we used a mirror to change the apparent number of items (Experiment 2), and tinted filters to change their apparent color (Experiment 3). In all three experiments, some chimpanzees were able to maximize their food rewards by making a choice based on the real properties of the stimuli in contrast to their manifest apparent properties. These results replicate the earlier findings for size illusions and extend them to additional situations involving illusory number and color. Control tests, together with findings from previous studies, ruled out lower-level explanations for the chimpanzees' performance. The findings thus support the hypothesis that chimpanzees are capable of making AR discriminations with a range of illusory stimuli.

A competitive nonverbal false belief task for children and apes.

A nonverbal false belief task was administered to children (mean age 5 years) and two great ape species: chimpanzees (Pan troglodytes) and bonobos (Pan paniscus). Because apes typically perform poorly in cooperative contexts, our task was competitive. Two versions were run: in both, a human competitor witnessed an experimenter hide a reward in one of two containers. When the competitor then left the room (version A) or turned around (version B), the experimenter switched the locations of the containers. The competitor returned and reached with effort, but unsuccessfully, towards the incorrect container. Children displayed an understanding of the competitor's false belief by correctly choosing the other container to find the reward. Apes did not. However, in version A (but not version B), apes looked more often at the unchosen container in false belief trials than in true belief control trials, possibly indicating some implicit or uncertain understanding that needs to be investigated further.

Can chimpanzees (Pan troglodytes) discriminate appearance from reality?

A milestone in human development is coming to recognize that how something looks is not necessarily how it is. We tested appearance-reality understanding in chimpanzees (Pan troglodytes) with a task requiring them to choose between a small grape and a big grape. The apparent relative size of the grapes was reversed using magnifying and minimizing lenses so that the truly bigger grape appeared to be the smaller one. Our Lens test involved a basic component adapted from standard procedures for children, as well as several components designed to rule out alternative explanations. There were large individual differences in performance, with some chimpanzees' responses suggesting they appreciated the appearance-reality distinction. In contrast, all chimpanzees failed a Reverse Contingency control test, indicating that those who passed the Lens test did not do so by learning a simple reverse contingency rule. Four-year-old children given an adapted version of the Lens test failed it while 4.5-year-olds passed. Our study constitutes the first direct investigation of appearance-reality understanding in chimpanzees and the first cross-species comparison of this capacity.

Chimpanzees (Pan troglodytes) know what can be seen from where.

Visual perspective taking research has established that chimpanzees recognize what others can or cannot see in the presence of occluding barriers. Less is known about chimpanzees' appreciation of what they themselves can or cannot see in similar situations. Additionally, it is unclear whether chimpanzees must rely on others' gaze cues to solve such tasks or whether they have a more general appreciation of what can be seen from where. Hence, we investigated chimpanzees' ability to anticipate what they would or would not be able to see from different visual perspectives. Food was hidden among arrays of open containers, with different containers providing visual access from unique viewing perspectives. Chimpanzees immediately adopted the correct perspective for each container type. Follow-up experiments showed that they were not simply moving to align themselves with visible openings. Our study thus suggests that chimpanzees have good visual perspective taking abilities with regard to themselves as well as others, and that both likely reflect a more general knowledge, at least implicit, of what can be seen from where.