Chimpanzees (Pan troglodytes) navigate to find hidden fruit in a virtual environment.

Almost all animals navigate their environment to find food, shelter, and mates. Spatial cognition of nonhuman primates in large-scale environments is notoriously difficult to study. Field research is ecologically valid, but controlling confounding variables can be difficult. Captive research enables experimental control, but space restrictions can limit generalizability. Virtual reality technology combines the best of both worlds by creating large-scale, controllable environments. We presented six chimpanzees with a seminaturalistic virtual environment, using a custom touch screen application. The chimpanzees exhibited signature behaviors reminiscent of real-life navigation: They learned to approach a landmark associated with the presence of fruit, improving efficiency over time; they located this landmark from novel starting locations and approached a different landmark when necessary. We conclude that virtual environments can allow for standardized testing with higher ecological validity than traditional tests in captivity and harbor great potential to contribute to longstanding questions in primate navigation, e.g., the use of landmarks, Euclidean maps, or spatial frames of reference.

Bo-NO-bouba-kiki: picture-word mapping but no spontaneous sound symbolic speech-shape mapping in a language trained bonobo.

Humans share the ability to intuitively map 'sharp' or 'round' pseudowords, such as 'bouba' versus 'kiki', to abstract edgy versus round shapes, respectively. This effect, known as sound symbolism, appears early in human development. The phylogenetic origin of this phenomenon, however, is unclear: are humans the only species capable of experiencing correspondences between speech sounds and shapes, or could similar effects be observed in other animals? Thus far, evidence from an implicit matching experiment failed to find evidence of this sound symbolic matching in great apes, suggesting its human uniqueness. However, explicit tests of sound symbolism have never been conducted with nonhuman great apes. In the present study, a language-competent bonobo completed a cross-modal matching-to-sample task in which he was asked to match spoken English words to pictures, as well as 'sharp' or 'round' pseudowords to shapes. Sound symbolic trials were interspersed among English words. The bonobo matched English words to pictures with high accuracy, but did not show any evidence of spontaneous sound symbolic matching. Our results suggest that speech exposure/comprehension alone cannot explain sound symbolism. This lends plausibility to the hypothesis that biological differences between human and nonhuman primates could account for the putative human specificity of this effect.

Chimpanzees (Pan troglodytes) show subtle signs of uncertainty when choices are more difficult.

Humans can tell when they find a task difficult. Subtle uncertainty behaviors like changes in motor speed and muscle tension precede and affect these experiences. Theories of animal metacognition likewise stress the importance of endogenous signals of uncertainty as cues that motivate metacognitive behaviors. However, while researchers have investigated second-order behaviors like information seeking and declining difficult trials in nonhuman animals, they have devoted little attention to the behaviors that express the cognitive conflict that gives rise to such behaviors in the first place. Here we explored whether three chimpanzees would, like humans, show hand wavering more when faced with more difficult choices in a touch screen transitive inference task. While accuracy was very high across all conditions, all chimpanzees wavered more frequently in trials that were objectively more difficult, demonstrating a signature behavior which accompanies experiences of difficulty in humans. This lends plausibility to the idea that feelings of uncertainty, like other emotions, can be studied in nonhuman animals. We propose to routinely assess uncertainty behaviors to inform models of procedural metacognition in nonhuman animals.

Using natural travel paths to infer and compare primate cognition in the wild.

Within comparative psychology, the evolution of animal cognition is typically studied either by comparing indirect measures of cognitive abilities (e.g., relative brain size) across many species or by conducting batteries of decision-making experiments among (typically) a few captive species. Here, we propose a third, complementary approach: inferring and comparing cognitive abilities through observational field records of natural information gradients and the associated variation in decision-making outcomes, using the ranging behavior of wild animals. To demonstrate the feasibility of our proposal, we present the results of a global survey assessing the availability of long-term ranging data sets from wild primates and the willingness of primatologists to share such data. We explore three ways in which such ranging data, with or without the associated behavioral and ecological data often collected by primatologists, might be used to infer and compare spatial cognition. Finally, we suggest how ecological complexity may be best incorporated into comparative analyses.

A Comparative Perspective on Three Primate Species' Responses to a Pictorial Emotional Stroop Task.

The Stroop effect describes interference in cognitive processing due to competing cognitive demands. Presenting emotionally laden stimuli creates similar Stroop-like effects that result from participants' attention being drawn to distractor stimuli. Here, we adapted the methods of a pictorial Stroop study for use with chimpanzees (N = 6), gorillas (N = 7), and Japanese macaques (N = 6). We tested all subjects via touchscreens following the same protocol. Ten of the 19 subjects passed pre-test training. Subjects who reached criterion were then tested on a standard color-interference Stroop test, which revealed differential accuracy in the primates' responses across conditions. Next, to test for an emotional Stroop effect, we presented subjects with photographs that were either positively valenced (a preferred food) or negatively valenced (snakes). In the emotional Stroop task, as predicted, the primates were less accurate in trials which presented emotionally laden stimuli as compared to control trials, but there were differences in the apes' and monkeys' response patterns. Furthermore, for both Stroop tests, while we found that subjects' accuracy rates were reduced by test stimuli, in contrast to previous research, we found no difference across trial types in the subjects' response latencies across conditions.

Chimpanzee Coordination and Potential Communication in a Two-touchscreen Turn-taking Game.

Recent years have seen a growing interest in the question of whether and how groups of nonhuman primates coordinate their behaviors for mutual benefit. On the one hand, it has been shown that chimpanzees in the wild and in captivity can solve various coordination problems. On the other hand, evidence of communication in the context of coordination problems is scarce. Here, we investigated how pairs of chimpanzees (Pan troglodytes) solved a problem of dynamically coordinating their actions for achieving a joint goal. We presented five pairs of chimpanzees with a turn-taking coordination game, where the task was to send a virtual target from one computer display to another using two touch-screens. During the joint practice of the game some subjects exhibited spontaneous gesturing. To address the question whether these gestures were produced to sustain coordination, we introduced a joint test condition in which we simulated a coordination break-down scenario: subjects appeared either unwilling or unable to return the target to their partner. The frequency of gesturing was significantly higher in these test trials than in the regular trials. Our results suggest that at least in some contexts chimpanzees can exhibit communicative behaviors to sustain coordination in joint action.

Sound symbolic congruency detection in humans but not in great apes.

Theories on the evolution of language highlight iconicity as one of the unique features of human language. One important manifestation of iconicity is sound symbolism, the intrinsic relationship between meaningless speech sounds and visual shapes, as exemplified by the famous correspondences between the pseudowords 'maluma' vs. 'takete' and abstract curved and angular shapes. Although sound symbolism has been studied extensively in humans including young children and infants, it has never been investigated in non-human primates lacking language. In the present study, we administered the classic "takete-maluma" paradigm in both humans (N = 24 and N = 31) and great apes (N = 8). In a forced choice matching task, humans but not great apes, showed crossmodal sound symbolic congruency effects, whereby effects were more pronounced for shape selections following round-sounding primes than following edgy-sounding primes. These results suggest that the ability to detect sound symbolic correspondences is the outcome of a phylogenetic process, whose underlying emerging mechanism may be relevant to symbolic ability more generally.

The word order of languages predicts native speakers' working memory.

The relationship between language and thought is controversial. One hypothesis is that language fosters habits of processing information that are retained even in non-linguistic domains. In left-branching (LB) languages, modifiers usually precede the head, and real-time sentence comprehension may more heavily rely on retaining initial information in working memory. Here we presented a battery of working memory and short-term memory tasks to adult native speakers of four LB and four right-branching (RB) languages from Africa, Asia and Europe. In working memory tasks, LB speakers were better than RB speakers at recalling initial stimuli, but worse at recalling final stimuli. Our results show that the practice of parsing sentences in specific directions due to the syntax and word order of our native language not only predicts the way we remember words, but also other non-linguistic stimuli.

Food cleaning in gorillas: Social learning is a possibility but not a necessity.

Food cleaning is widespread in the animal kingdom, and a recent report confirmed that (amongst other behaviours) wild western lowland gorillas also show food cleaning. The authors of this report conclude that this behaviour, based on its distribution patterns, constitutes a potential candidate for culture. While different conceptualisations of culture exist, some more and some less reliant on behavioural form copying, all of them assign a special role to social learning processes in explaining potentially cultural behaviours. Here we report the results of an experiment that tested to what extent food cleaning behaviour in a group of captive Western lowland gorillas (Gorilla gorilla gorilla) relies on social learning processes. Subjects were provided with clean and dirty apples. When they were provided with dirty apples, all subjects showed evidence of food cleaning in at least 75% of trials. Preferred cleaning techniques differed between individuals, four out of five of subjects expressed a behaviour analogous to that reported in wild conspecifics. Given this occurrence of food cleaning in a culturally unconnected population of gorillas, we conclude that social learning is unlikely to play a central role in the emergence of the food cleaning behavioural form in Western lowland gorillas; instead, placing a greater emphasis on individual learning of food cleaning's behavioural form.

Information seeking about tool properties in great apes.

Evidence suggests that great apes engage in metacognitive information seeking for food items. To support the claim that a domain-general cognitive process underlies ape metacognition one needs to show that selective information seeking extends to non-food items. In this study, chimpanzees (Pan troglodytes) and orangutans (Pongo abelii) either had to determine the location of a desired food item or a property of a non-food item (length of a tool). We manipulated whether subjects received prior information about the item's location or property. During the test, subjects had the opportunity to seek the respective information. Results show that apes engaged in more information seeking when they had no prior knowledge. Importantly, this selective pattern of information seeking applied to food as well as to tools.

How chimpanzees (Pan troglodytes) perform in a modified emotional Stroop task.

The emotional Stroop task is an experimental paradigm developed to study the relationship between emotion and cognition. Human participants required to identify the color of words typically respond more slowly to negative than to neutral words (emotional Stroop effect). Here we investigated whether chimpanzees (Pan troglodytes) would show a comparable effect. Using a touch screen, eight chimpanzees were trained to choose between two simultaneously presented stimuli based on color (two identical images with differently colored frames). In Experiment 1, the images within the color frames were shapes that were either of the same color as the surrounding frame or of the alternative color. Subjects made fewer errors and responded faster when shapes were of the same color as the frame surrounding them than when they were not, evidencing that embedded images affected target selection. Experiment 2, a modified version of the emotional Stroop task, presented subjects with four different categories of novel images: three categories of pictures of humans (veterinarian, caretaker, and stranger), and control stimuli showing a white square. Because visits by the veterinarian that include anaesthetization can be stressful for subjects, we expected impaired performance in trials presenting images of the veterinarian. For the first session, we found correct responses to be indeed slower in trials of this category. This effect was more pronounced for subjects whose last anaesthetization experience was more recent, indicating that emotional valence caused the slowdown. We propose our modified emotional Stroop task as a simple method to explore which emotional stimuli affect cognitive performance in nonhuman primates.

Cross-cultural evidence of value structures and priorities in childhood.

We broaden the developmental focus of the theory of universals in basic human values (Schwartz, 1992, Advances in Experimental Social Psychology) by presenting supportive evidence on children's values from six countries: Germany, Italy, Poland, Bulgaria, the United States, and New Zealand. 3,088 7-11-year-old children completed the Picture-Based Value Survey for Children (PBVS-C, Döring et al., 2010, J. Pers. Assess., 92, 439). Grade 5 children also completed the Portrait Values Questionnaire (PVQ, Schwartz, 2003, A proposal for measuring value orientations across nations. Chapter 7 in the Questionnaire Development Package of the European Social Survey). Findings reveal that the broad value structures, sex differences in value priorities and pan-cultural value hierarchies typical of adults have already taken form at this early age. We discuss the conceptual implications of these findings for the new field of children's basic values by embedding them in the recent developmental literature.

Food washing and placer mining in captive great apes.

Sweet potato washing and wheat placer mining in Japanese macaques (Macaca fuscata) are among the most well known examples of local traditions in non-human animals. The functions of these behaviors and the mechanisms of acquisition and spread of these behaviors have been debated frequently. Prompted by animal caretaker reports that great apes [chimpanzees (Pan troglodytes), bonobos (Pan paniscus), gorillas (Gorilla gorilla), and orangutans (Pongo abelii)] at Leipzig Zoo occasionally wash their food, we conducted a study of food washing behaviors that consisted of two parts. In the first part we assessed the current distribution of the behavior on the basis of caretaker reports. In the second (experimental) part, we provided subjects individually with a water basin and two types of food (apples and cereal) that was either clean or covered/mixed with sand. We found that subjects of all species (except gorillas) placed apples in the water before consumption, and that they did so more often when the apples were dirty than when they were clean. Several chimpanzees and orangutans also engaged in behaviors resembling wheat placer mining.