Christoph J. Völter

Repeated innovation in great apes

Innovation has been defined as a solution to a novel problem or a novel solution to an old problem. The second part of this definition requires the inhibition of previously learnt solution strategies before a novel solution can be found. Therefore, inventing novel solutions for an old problem is considered to be particularly difficult. We investigated the ability of great apes to produce multiple new solutions to a task after each of those solutions became obsolete. We presented all four nonhuman great ape species with a task consisting of extracting a food reward from a puzzle box. Initially, the task could be solved in three different ways that varied in difficulty. After subjects discovered the first solution, we allowed them to use it for some trials and then it became obsolete. If the apes could overcome their initial response and find the next solution, we again allowed them to use it for some time and once again it became obsolete. The final step consisted of finding the third solution to secure the food reward. We found that all species except orang-utans, Pongo abelii, were able to solve all versions of the problem. Furthermore, they overcame the obsolete techniques quickly and efficiently, indicating high degrees of behavioural flexibility and inhibitory control. In contrast to previous research on social learning, our results suggest that great apes are not conservative and adjust their behaviour flexibly when the physical constraints of a task change.

Problem solving in great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo abelii): the effect of visual feedback

What kind of information animals use when solving problems is a controversial topic. Previous research suggests that, in some situations, great apes prefer to use causally relevant cues over arbitrary ones. To further examine to what extent great apes are able to use information about causal relations, we presented three different puzzle box problems to the four nonhuman great ape species. Of primary interest here was a comparison between one group of apes that received visual access to the functional mechanisms of the puzzle boxes and one group that did not. Apes’ performance in the first two, less complex puzzle boxes revealed that they are able to solve such problems by means of trial-and-error learning, requiring no information about the causal structure of the problem. However, visual inspection of the functional mechanisms of the puzzle boxes reduced the amount of time needed to solve the problems. In the case of the most complex problem, which required the use of a crank, visual feedback about what happened when the handle of the crank was turned was necessary for the apes to solve the task. Once the solution was acquired, however, visual feedback was no longer required. We conclude that visual feedback about the consequences of their actions helps great apes to solve complex problems. As the crank task matches the basic requirements of vertical string pulling in birds, the present results are discussed in light of recent findings with corvids.

Younger apes and human children plan their moves in a maze task

Planning defined as the predetermination of a sequence of actions towards some goal is crucial for complex problem solving. To shed light on the evolution of executive functions, we investigated the ontogenetic and phylogenetic origins of planning. Therefore, we presented all four great apes species (N=12) as well as 4- and 5-year-old human preschoolers (N=24) with a vertical maze task. To gain a reward placed on the uppermost level of the maze, subjects had to move the reward to the bottom through open gaps situated at each level of the maze. In total, there were ten gaps located over three of the mazes levels, and free passage through these gaps could be flexibly blocked using multiple traps. Due to the decision tree design of the maze, the subjects had to plan their actions depending on the trap configuration up to two steps ahead to successfully retrieve the reward. We found that (1) our measure of planning was negatively correlated with age in nonhuman apes, (2) younger apes as well as 5-year-old children planned their moves up to two steps ahead whereas 4-year-olds were limited to plan one step ahead, and (3) similar performance but different underlying limitations between apes and children. Namely, while all species of nonhuman apes were limited by a lack of motor control, human children exhibited a shortage in shifting their attention across a sequence of subgoals.

Great apes and children infer causal relations from patterns of variation and covariation

We investigated whether nonhuman great apes (N=23), 2.5-year-old (N=20), and 3-year-old children (N=40) infer causal relations from patterns of variation and covariation by adapting the blicket detector paradigm for apes. We presented chimpanzees (Pan troglodytes), bonobos (Pan paniscus), orangutans (Pongo abelii), gorillas (Gorilla gorilla), and children (Homo sapiens) with a novel reward dispenser, the blicket detector. The detector was activated by inserting specific (yet randomly determined) objects, the so-called blickets. Once activated a reward was released, accompanied by lights and a short tone. Participants were shown different patterns of variation and covariation between two different objects and the activation of the detector. When subsequently choosing between one of the two objects to activate the detector on their own all species, except gorillas (who failed the training), took these patterns of correlation into account. In particular, apes and 2.5-year-old children ignored objects whose effect on the detector completely depended on the presence of another object. Follow-up experiments explored whether the apes and children were also able to re-evaluate evidence retrospectively. Only children (3-year-olds in particular) were able to make such retrospective inferences about causal structures from observing the effects of the experimenters actions. Apes succeeded here only when they observed the effects of their own interventions. Together, this study provides evidence that apes, like young children, accurately infer causal structures from patterns of (co)variation and that they use this information to inform their own interventions.

Comparative psychometrics: establishing what differs is central to understanding what evolves

Cognitive abilities cannot be measured directly. What we can measure is individual variation in task performance. In this paper, we first make the case for why we should be interested in mapping individual differences in task performance onto particular cognitive abilities: we suggest that it is crucial for examining the causes and consequences of variation both within and between species. As a case study, we examine whether multiple measures of inhibitory control for non-human animals do indeed produce correlated task performance; however, no clear pattern emerges that would support the notion of a common cognitive ability underpinning individual differences in performance. We advocate a psychometric approach involving a three-step programme to make theoretical and empirical progress: first, we need tasks that reveal signature limits in performance. Second, we need to assess the reliability of individual differences in task performance. Third, multi-trait multi-method test batteries will be instrumental in validating cognitive abilities. Together, these steps will help us to establish what varies between individuals that could impact their fitness and ultimately shape the course of the evolution of animal minds. Finally, we propose executive functions, including working memory, inhibitory control and attentional shifting, as a sensible starting point for this endeavour. This article is part of the theme issue ‘Causes and consequences of individual differences in cognitive abilities’.

Cooperative problem solving in giant otters (Pteronura brasiliensis) and Asian small-clawed otters (Aonyx cinerea)

Cooperative problem solving has gained a lot of attention over the past two decades, but the range of species studied is still small. This limits the possibility of understanding the evolution of the socio-cognitive underpinnings of cooperation. Lutrinae show significant variations in socio-ecology, but their cognitive abilities are not well studied. In the first experimental study of otter social cognition, we presented two species—giant otters and Asian small-clawed otters—with a cooperative problem-solving task. The loose string task requires two individuals to simultaneously pull on either end of a rope in order to access food. This task has been used with a larger number of species (for the most part primates and birds) and thus allows for wider cross-species comparison. We found no differences in performance between species. Both giant otters and Asian small-clawed otters were able to solve the task successfully when the coordination requirements were minimal. However, when the temporal coordination demands were increased, performance decreased either due to a lack of understanding of the role of a partner or due to difficulty inhibiting action. In conclusion, two species of otters show some ability to cooperate, quite similar to most other species presented with the same task. However, to draw further conclusions and more nuanced comparisons between the two otter species, further studies with varied methodologies will be necessary.

Social manipulation in nonhuman primates: Cognitive and motivational determinants.

HighlightsWe classify the experimental literature on primate social interactions along 2 axes.A cognitive axis includes problem‐solving, social tool use, and communication.A motivational axis includes self‐ and other‐regarding preferences.Great apes seem particularly adept among primates at manipulating conspecifics.The evidence for other‐regarding motives supporting social interactions is inconsistent. Abstract Social interactions are the result of individuals’ cooperative and competitive tendencies expressed over an extended period of time. Although social manipulation, i.e., using another individual to achieve one’s own goals, is a crucial aspect of social interactions, there has been no comprehensive attempt to differentiate its various types and to map its cognitive and motivational determinants. For this purpose, we survey in this article the experimental literature on social interactions in nonhuman primates. We take social manipulation, illustrated by a case study with orangutans (Pongo abelii), as our starting point and move in two directions. First, we will focus on a flexibility/sociality axis that includes technical problem solving, social tool‐use and communication. Second, we will focus on a motivational/prosociality axis that includes exploitation, cooperation, and helping. Combined, the two axes offer a way to capture a broad range of social interactions performed by human and nonhuman primates.

Intuitive optics : what great apes infer from mirrors and shadows

There is ongoing debate about the extent to which nonhuman animals, like humans, can go beyond first-order perceptual information to abstract structural information from their environment. To provide more empirical evidence regarding this question, we examined what type of information great apes (chimpanzees, bonobos, and orangutans) gain from optical effects such as shadows and mirror images. In an initial experiment, we investigated whether apes would use mirror images and shadows to locate hidden food. We found that all examined ape species used these cues to find the food. Follow-up experiments showed that apes neither confused these optical effects with the food rewards nor did they merely associate cues with food. First, naïve chimpanzees used the shadow of the hidden food to locate it but they did not learn within the same number of trials to use a perceptually similar rubber patch as indicator of the hidden food reward. Second, apes made use of the mirror images to estimate the distance of the hidden food from their own body. Depending on the distance, apes either pointed into the direction of the food or tried to access the hidden food directly. Third, apes showed some sensitivity to the geometrical relation between mirror orientation and mirrored objects when searching hidden food. Fourth, apes tended to interpret mirror images and pictures of these mirror images differently depending on their prior knowledge. Together, these findings suggest that apes are sensitive to the optical relation between mirror images and shadows and their physical referents.