Keeping track of time: evidence for episodic-like memory in great apes

Episodic memory, as defined by Tulving, can be described in terms of behavioural elements (what, where and when information) but it is also accompained by an awareness of one’s past (chronesthesia) and a subjective conscious experience (autonoetic awareness). Recent experiments have shown that corvids and rodents recall the where, what and when of an event. This capability has been called episodic-like memory because it only fulfils the behavioural criteria for episodic memory. We tested seven chimpanzees, three orangutans and two bonobos of various ages by adapting two paradigms, originally developed by Clayton and colleagues to test scrub jays. In Experiment 1, subjects were fed preferred but perishable food (frozen juice) and less preferred but non-perishable food (grape). After the food items were hidden, subjects could choose one of them either after 5 min or 1 h. The frozen juice was still available after 5 min but melted after 1 h and became unobtainable. Apes chose the frozen juice significantly more after 5 min and the grape after 1 h. In Experiment 2, subjects faced two baiting events happening at different times, yet they formed an integrated memory for the location and time of the baiting event for particular food items. We also included a memory task that required no temporal encoding. Our results showed that apes remember in an integrated fashion what, where and when (i.e., how long ago) an event happened; that is, apes distinguished between different events in which the same food items were hidden in different places at different times. The temporal control of their choices was not dependent on the familiarity of the platforms where the food was hidden. Chimpanzees’ and bonobos’ performance in the temporal encoding task was age-dependent, following an inverted U-shaped distribution. The age had no effect on the performance of the subjects in the task that required no temporal encoding.     

Natural reference: A phylo‐ and ontogenetic perspective on the comprehension of iconic gestures and vocalizations

The recognition of iconic correspondence between signal and referent has been argued to bootstrap the acquisition and emergence of language. Here, we study the ontogeny, and to some extent the phylogeny, of the ability to spontaneously relate iconic signals, gestures, and/or vocalizations, to previous experience. Children at 18, 24, and 36 months of age (N = 216) and great apes (N = 13) interacted with two apparatuses, each comprising a distinct action and sound. Subsequently, an experimenter mimicked either the action, the sound, or both in combination to refer to one of the apparatuses. Experiments 1 and 2 found no spontaneous comprehension in great apes and in 18‐month‐old children. At 24 months of age, children were successful with a composite vocalization‐gesture signal but not with either vocalization or gesture alone. At 36 months, children succeeded both with a composite vocalization‐gesture signal and with gesture alone, but not with vocalization alone. In general, gestures were understood better compared to vocalizations. Experiment 4 showed that gestures were understood irrespective of how children learned about the corresponding action (through observation or self‐experience). This pattern of results demonstrates that iconic signals can be a powerful way to establish reference in the absence of language, but they are not trivial for children to comprehend and not all iconic signals are created equal.     

Orangutans (Pongo pygmaeus) and bonobos (Pan paniscus) point to inform a human about the location of a tool

Although pointing is not part of great apes’ natural gestural repertoire, they can learn to point to food, in order to request it. To assess the flexibility with which they can use this gesture, one can vary the potential referent of the point. In two previous studies, three orangutans (two of them human-reared) have shown the ability to point to the location of a tool which a human experimenter needed in order to give them food. Here, we tested six orangutans and five bonobos using a set-up in which our subjects had to guide a human experimenter to the hiding place of a fork which was needed in order to retrieve a piece of food for the subject out of a vertical tube. We further examined the potential role of a competitive/deceptive context by varying the identity of the person responsible for hiding the tool. In addition, we implemented three different control conditions in which an object was hidden but it was not necessary to indicate its location to get the food. We found that the majority of subjects spontaneously guided the experimenter to the hiding place of the fork by pointing to it when it was necessary and they did so significantly less in control conditions. We did not find an effect of the person hiding the fork. Our results show that mother-reared orangutans and bonobos are able to point to inform a human about the location of an object that the human needs to procure food for the subject and that they can take into account whether it is relevant or not to do so.

Comparing dogs and great apes in their ability to visually track object transpositions

Knowing that objects continue to exist after disappearing from sight and tracking invisible object displacements are two basic elements of spatial cognition. The current study compares dogs and apes in an invisible transposition task. Food was hidden under one of two cups in full view of the subject. After that both cups were displaced, systematically varying two main factors, whether cups were crossed during displacement and whether the cups were substituted by the other cup or instead cups were moved to new locations. While the apes were successful in all conditions, the dogs had a strong preference to approach the location where they last saw the reward, especially if this location remained filled. In addition, dogs seem to have especial difficulties to track the reward when both containers crossed their path during displacement. These results confirm the substantial difference that exists between great apes and dogs with regard to mental representation abilities required to track the invisible displacements of objects.     

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.

Prevalence of renewal of problem behavior during context changes

Renewal is defined as the reemergence of a previously eliminated behavior following a context change. Determining the prevalence of this effect in clinical practice would allow clinicians to better anticipate the reemergence of problem behavior, such as when a patient is discharged from a treatment facility to return to their home. The current consecutive, case-series analysis determined the prevalence and magnitude of renewal when implementing behavioral treatments for problem behavior. Across 182 context changes, renewal was observed 77 times (42.3%). In the first session following the context change, problem behavior rates increased by a factor of 3 and then decreased across successive sessions. These results indicated that renewal effects may be common, but are also transient and return to rates observed before context changes.     

Inferences by exclusion in the great apes: the effect of age and species

This study investigated the ability of chimpanzees, gorillas, orangutans, and bonobos to make inferences by exclusion using the procedure pioneered by Premack and Premack (Cognition 50:347–362, 1994) with chimpanzees. Thirty apes were presented with two different food items (banana vs. grape) on a platform and covered with identical containers. One of the items was removed from the container and placed between the two containers so that subjects could see it. After discarding this item, subjects could select between the two containers. In Experiment 1, apes preferentially selected the container that held the item that the experimenter had not discarded, especially if subjects saw the experimenter remove the item from the container (but without seeing the container empty). Experiment 3 in which the food was removed from one of the containers behind a barrier confirmed these results. In contrast, subjects performed at chance levels when a stimulus (colored plastic chip: Exp. 1; food item: Exp. 2 and Exp. 3) designated the item that had been removed. These results indicated that apes made inferences, not just learned to use a discriminative cue to avoid the empty container. Apes perceived and treated the item discarded by the experimenter as if it were the very one that had been hidden under the container. Results suggested a positive relationship between age and inferential ability independent of memory ability but no species differences.This contribution is part of the special issue “Animal Logics” (Watanabe and Huber 2006).