Chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) quantify split solid objects

Recent research suggests that gorillas’ and orangutans’ object representations survive cohesion violations (e.g., a split of a solid object into two halves), but that their processing of quantities may be affected by them. We assessed chimpanzees’ (Pan troglodytes) and bonobos’ (Pan paniscus) reactions to various fission events in the same series of action tasks modelled after infant studies previously run on gorillas and orangutans (Cacchione and Call in Cognition 116:193–203, 2010b). Results showed that all four non-human great ape species managed to quantify split objects but that their performance varied as a function of the non-cohesiveness produced in the splitting event. Spatial ambiguity and shape invariance had the greatest impact on apes’ ability to represent and quantify objects. Further, we observed species differences with gorillas performing lower than other species. Finally, we detected a substantial age effect, with ape infants below 6 years of age being outperformed by both juvenile/adolescent and adult apes.     

Great apes' (Pan troglodytes, Pan paniscus, Gorilla gorilla, Pongo pygmaeus) understanding of tool functional properties after limited experience

Primates' understanding of tool functionality has been investigated extensively using a paradigm in which subjects are presented with a tool that they must use to obtain an out-of-reach reward. After being given experience on an initial problem, monkeys can transfer their skill to tools of different shapes while ignoring irrelevant tool changes (e.g., color). In contrast, monkeys without initial training perform poorly on the same tasks. Compared to most monkeys, great apes show a clear propensity for tool using and may not require as much experience to succeed on tool functionality tasks. We investigated this question by presenting 171 apes (Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus) with several tool-use problems without giving them initial training or familiarizing them with the test materials. Apes succeeded without experience, but only on problems based on basic properties such as the reward being supported by an object. However, only minimal experience was sufficient to allow them to quickly improve their performance on more complex problems in which the reward was not in contact with the tool.     

Object permanence in orangutans (Pongo pygmaeus), chimpanzees (Pan troglodytes), and children (Homo sapiens).

Juvenile and adult orangutans (n = 5; Pongo pygmaeus), chimpanzees (n = 7; Pan troglodytes), and 19- and 26-month-old children (n = 24; Homo sapiens) received visible and invisible displacements. Three containers were presented forming a straight line, and a small box was used to displace a reward under them. Subjects received 3 types of displacement: single (the box visited 1 container), double adjacent (the box visited 2 contiguous containers), and double nonadjacent (the box visited 2 noncontiguous containers). All species performed at comparable levels, solving all problems except the invisible nonadjacent displacements. Visible displacements were easier than invisible, and single were easier than double displacements. In a 2nd experiment, subjects saw the baiting of either 2 adjacent or 2 nonadjacent containers with no displacements. All species selected the empty container more often when the baited containers were nonadjacent than when they were adjacent. It is hypothesized that a response bias and inhibition problem were responsible for the poor performance in nonadjacent displacements.     

Chimpanzees (Pan troglodytes) remember the location of a hidden food item after altering their orientation to a spatial array

Two chimpanzees (Pan troglodytes) had a direct view of an experimenter placing a food item beneath one of several cups within a horizontal spatial array. The chimpanzees then were required to move around the spatial array, shifting their orientation to the array by 180 degrees . Both chimpanzees remembered the location of the food item. In the next experiment, a visual barrier was placed between the chimpanzees and the spatial array after the food item had been hidden to prevent visual tracking of the location of the object during the chimpanzees' movement. One chimpanzee remembered the location of the hidden item in this variation. These results demonstrate another capacity for spatial memory in this species that complements data indicating chimpanzee spatial memory for invisible displacements, array rotations, and array transpositions.     

Piagetian liquid conservation in the great apes (Pan paniscus, Pan troglodytes, and Pongo pygmaeus)

An understanding of Piagetian liquid conservation was investigated in 4 bonobos (Pan paniscus), 5 chimpanzees (Pan troglodytes), and 5 orangutans (Pongo pygmaeus). The apes were tested in the ability to track the larger of 2 quantities of juice that had undergone various kinds of transformations. The accuracy of the apes' judgment depended on the shape or number of containers into which the larger quantity was transferred. The apes made their choice mainly on the basis of visual estimation but showed modest success when the quantities were occluded. The results suggest that the apes rely to a greater extent on visual information, although they might have some appreciation of the constancy of liquid quantities.     

Strategies used to combine seriated cups by chimpanzees (Pan troglodytes), bonobos (Pan paniscus), and capuchins (Cebus apella).

The authors investigated strategies used to combine seriated cups by apes (Pan troglodytes and P. paniscus) and monkeys (Cebus apella) using a protocol reported in P. M. Greenfield, K. Nelson, and E. Saltzman's (1972) study with children. It was hypothesized that apes would exhibit more hierarchical combinations of cups than monkeys, given apes' language capacity, and that apes would seriate the cups more efficiently than monkeys. As predicted, apes made many structures with the cups using a variety of strategies, and monkeys rarely combined the cups. After a training phase to orient monkeys to the task, the 2 genera did not differ in the strategies used to combine the cups or in efficiency in seriating the cups. Success in this task suggests that sensorimotor versions of hierarchically organized combinatorial activity are well within apes' and monkeys' abilities.     

Tubes,tables and traps: great apes solve two functionally equivalent trap tasks but show no evidence of transfer across tasks

Previous studies on tool using have shown that presenting subjects with certain modifications in the experimental setup can substantially improve their performance. However, procedural modifications (e.g. trap table task) may not only remove task constraints but also simplify the problem conceptually. The goal of this study was to design a variation of the trap-table that was functionally equivalent to the trap-tube task. In this new task, the subjects had to decide where to insert the tool and in which direction the reward should be pushed. We also administered a trap-tube task that allowed animals to push or rake the reward with the tool to compare the subjects' performance on both tasks. We used a larger sample of subjects than in previous studies and from all the four species of great apes (Gorilla gorilla, Pan troglodytes, Pan paniscus, and Pongo pygmaeus). The results showed that apes performed better in the trap-platform task than in the trap-tube task. Subjects solved the tube task faster than in previous studies and they also preferred to rake in rather than to push the reward out. There was no correlation in the level of performance between both tasks, and no indication of interspecies differences. These data are consistent with the idea that apes may possess some specific causal knowledge of traps but may lack the ability to establish analogical relations between functional equivalent tasks.     

How young children and chimpanzees (Pan troglodytes) perceive objects in a 2D display: putting an assumption to the test

Object recognition research is typically conducted using 2D stimuli in lieu of 3D objects. This study investigated the amount and complexity of knowledge gained from 2D stimuli in adult chimpanzees (Pan troglodytes) and young children (aged 3 and 4 years) using a titrated series of cross-dimensional search tasks. Results indicate that 3-year-old children utilize a response rule guided by local features to solve cross-dimensional tasks. Four-year-old toddlers and adult chimpanzees use information about object form and compositional structure from a 2D image to guide their search in three dimensions. Findings have specific implications to research conducted in object recognition/perception and broad relevance to all areas of research and daily living that incorporate 2D displays.     

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) perform on the reversed contingency task: the effects of food quantity and food visibility

S. T. Boysen and G. G. Berntson (1995) found that chimpanzees performed poorly on a reversed contingency task in which they had to point to the smaller of 2 food quantities to acquire the larger quantity. The authors compared the performance of 4 great ape species (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) on the reversed contingency task while manipulating food quantity (0-4 or 1-4) and food visibility (visible pairs or covered pairs). Results showed no systematic species differences but large individual differences. Some individuals of each species were able to solve the reversed contingency task. Both quantity and visibility of the food items had a significant effect on performance. Subjects performed better when the disparity between quantities was smaller and the quantities were not directly visible.     

Maintenance of self-imposed delay of gratification by four chimpanzees (Pan troglodytes) and an orangutan (Pongo pygmaeus)

Delay maintenance, which is the continuance over time of the choice to forgo an immediate, less preferred reward for a future, more preferred reward, was examined in 4 chimpanzees (Pan troglodytes) and 1 orangutan (Pongo pygmaeus). In the 1st experiment, the apes were presented with 20 chocolate pieces that were placed, one at a time, into a bowl that was within their reach. The apes could consume the available chocolate pieces at any time during a trial, but no additional pieces would be given. The total length of time taken to place the 20 items into the bowl ranged from 60 s to 180 s. All 5 apes delayed gratification on a majority of trials until all 20 chocolate pieces were presented. Unlike in most experiments with human children using this test situation, attention by the apes to the reward was not detrimental to delay maintenance. In a 2nd experiment with the chimpanzees only, 4 foods of differing incentive value were presented in different trials in the same manner as in Experiment 1. The chimpanzees were highly successful in obtaining all food pieces, and there was no difference in performance as a function of food type.     

Referential understanding of videos in chimpanzees (Pan troglodytes), orangutans (Pongo pygmaeus), and children (Homo sapiens)

Performance on identical search tasks based on cues directly perceived or indirectly perceived through video were compared among a group of 4 adult chimpanzees (Pan troglodytes), a group of 2 adult orangutans (Pongo pygmaeus), and a group of 36 children (between 2 and 3 years of age). Children comprehended directly perceived cues but had difficulty with video cues. In contrast, chimpanzees and 1 orangutan were successful in using video to guide their search for a hidden object. Two follow-up studies with 3-year-old children demonstrated the importance of more distinct perceptual and verbal cues in aiding children's understanding of video as referring to real-world events.     

Apes' and children's understanding of cooperative and competitive motives in a communicative situation

Chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) (Study 1) and 18- and 24-month-old human children (Study 2) participated in a novel communicative task. A human experimenter (E) hid food or a toy in one of two opaque containers before gesturing towards the reward's location in one of two ways. In the Informing condition, she attempted to help the subject find the hidden object by simply pointing to the correct container. In the Prohibiting condition, E held out her arm toward the correct container (palm out) and told the subject firmly 'Don't take this one.' As in previous studies, the apes were at chance in the Informing condition. However, they were above chance in the new Prohibiting condition. Human 18-month-olds showed this same pattern of results, whereas 24-month-olds showed the opposite pattern: they were better in the Informing condition than in the Prohibiting condition. In our interpretation, success in the Prohibiting condition requires subjects to understand E's goal toward them and their behavior, and then to make an inference (she would only prohibit if there were something good in there). Success in the Informing condition requires subjects to understand a cooperative communicative motive - which apparently apes and young infants find difficult.     

Spatial construction skills of chimpanzees (Pan troglodytes) and young human children (Homo sapiens sapiens)

Spatial construction tasks are basic tests of visual‐spatial processing. Two studies have assessed spatial construction skills in chimpanzees (Pan troglodytes) and young children (Homo sapiens sapiens) with a block modelling task. Study 1a subjects were three young chimpanzees and five adult chimpanzees. Study 1b subjects were 30 human children belonging to five age groups (24, 30, 36, 42, 48 months). Subjects were given three model constructions to reproduce: Line, Cross‐Stack and Arch, which differed in type and number of spatial relations and dimensions, but required comparable configurational understanding. Subjects’ constructions were rated for accuracy. Our results show that: (1) chimpanzees are relatively advanced in constructing in the vertical dimension; (2) Among chimpanzees only adults make accurate copies of constructions; (3) Chimpanzees do not develop in the direction of constructing in two dimensions as human children do starting from age 30 months. The pattern of development of construction skills in chimpanzees partially diverges from that of human children and indicates that spatial analysis and spatial representation are partially different in the two species.     

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, Gorilla gorilla) perform on the reversed reward contingency task II: transfer to new quantities, long-term retention, and the impact of quantity ratios

We tested 6 chimpanzees (Pan troglodytes), 3 orangutans (Pongo pygmaeus), 4 bonobos (Pan paniscus), and 2 gorillas (Gorilla gorilla) in the reversed reward contingency task. Individuals were presented with pairs of quantities ranging between 0 and 6 food items. Prior to testing, some experienced apes had solved this task using 2 quantities while others were totally na?ve. Experienced apes transferred their ability to multiple-novel pairs after 6 to 19 months had elapsed since their initial testing. Two out of 6 na?ve apes (1 chimpanzee, 1 bonobo) solved the task--a proportion comparable to that of a previous study using 2 pairs of quantities. Their acquisition speed was also comparable to the successful subjects from that study. The ratio between quantities explained a large portion of the variance but affected na?ve and experienced individuals differently. For smaller ratios, na?ve individuals were well below 50% correct and experienced ones were well above 50%, yet both groups tended to converge toward 50% for larger ratios. Thus, some apes require no procedural modifications to overcome their strong bias for selecting the larger of 2 quantities.     

Comparative assessment of handedness for a coordinated bimanual task in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus)

Hand preferences for a coordinated bimanual task were assessed in a sample of 31 captive gorillas (Gorilla gorilla) and 19 captive orangutans (Pongo pygmaeus) and were compared with chimpanzee (Pan troglodytes) hand preferences in subjects that were matched on the basis of age, sex, and rearing history. The task required that the apes remove food from the inside edges of a symmetrical polyvinyl chloride pipe presented to them in their home cages. The results indicate significant species differences with chimpanzees showing population-level right-handedness and orangutans showing population-level left-handedness. The gorillas showed a nonsignificant trend toward right-handedness. The results are discussed in terms of possible ecological or biomechanical factors that may influence hand preferences in different ape species.     

Making inferences about the location of hidden food: social dog, causal ape

Domestic dogs (Canis familiaris) and great apes from the genus Pan were tested on a series of object choice tasks. In each task, the location of hidden food was indicated for subjects by some kind of communicative, behavioral, or physical cue. On the basis of differences in the ecologies of these 2 genera, as well as on previous research, the authors hypothesized that dogs should be especially skillful in using human communicative cues such as the pointing gesture, whereas apes should be especially skillful in using physical, causal cues such as food in a cup making noise when it is shaken. The overall pattern of performance by the 2 genera strongly supported this social-dog, causal-ape hypothesis. This result is discussed in terms of apes' adaptations for complex, extractive foraging and dogs' adaptations, during the domestication process, for cooperative communication with humans.     

What meaning means for same and different: Analogical reasoning in humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta)

Thus far, language- and token-trained apes (e.g., D. Premack, 1976; R. K. R. Thompson, D. L. Oden, & S. T. Boysen, 1997) have provided the best evidence that nonhuman animals can solve, complete, and construct analogies, thus implicating symbolic representation as the mechanism enabling the phenomenon. In this study, the authors examined the role of stimulus meaning in the analogical reasoning abilities of three different primate species. Humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta) completed the same relational matching-to-sample (RMTS) tasks with both meaningful and nonmeaningful stimuli. This discrimination of relations-between-relations serves as the basis for analogical reasoning. Meaningfulness facilitated the acquisition of analogical matching for human participants, whereas individual differences among the chimpanzees suggest that meaning can either enable or hinder their ability to complete analogies. Rhesus monkeys did not succeed in the RMTS task regardless of stimulus meaning, suggesting that their ability to reason analogically, if present at all, may be dependent on a dimension other than the representational value of stimuli.     

Object permanence in common marmosets (Callithrix jacchus)

A series of 9 search tasks corresponding to the Piagetian Stages 3-6 of object permanence were administered to 11 common marmosets (Callithrix jacchus). Success rates varied strongly among tasks and marmosets, but the performances of most subjects were above chance level on the majority of tasks of visible and invisible displacements. Although up to 24 trials were administered in the tests, subjects did not improve their performance across trials. Errors were due to preferences for specific locations or boxes, simple search strategies, and attentional deficits. The performances of at least 2 subjects that achieved very high scores up to the successive invisible displacement task suggest that this species is able to represent the existence and the movements of unperceived objects.     

Chimpanzees (Pan troglodytes) respond to nonvisible sets after one-by-one addition and removal of items

Two chimpanzees (Pan troglodytes) made numerousness judgments of nonvisible sets of items. In Experiment 1, 1-10 items were dropped 1 at a time into an opaque cup, and then an additional 1-10 items were dropped 1 at a time into another opaque cup. The chimpanzees' performance levels were high and were more dependent on factors indicative of an analogue-magnitude mechanism for representation of set size than on an object file mechanism. In Experiment 2, a 3rd visible set was made available after the sequential presentation of the first 2 sets. The chimpanzees again performed at high levels in selecting the largest of the 3 sets. In Experiment 3, 1 of the 2 initially presented sets was reduced in number by the sequential removal of 1, 2, or 3 items. Both chimpanzees performed above chance levels for the removal of 1, but not more than 1, item.     

Development of combinatory manipulation in chimpanzee infants (Pan troglodytes)

I made systematic observations of three infant chimpanzees aged 2–4 years, who participated in a series of diagnostic tests of combinatory manipulation. The tasks were stacking blocks, seriating nesting cups, and inserting an object into the corresponding hole in a plate or a box. These tasks were originally devised for developmental diagnosis of human infants. The chimpanzee infants displayed combinatory manipulation comparable to that of 1-year-old human infants. Common motor characteristics were observed across the tasks, namely "repetition" of actions, "adjustment" of actions, "reversal" of actions, and "shifts" of attention. Humans and chimpanzees share these actions when manipulating multiple objects to complete a task. Repetition, adjustment, and reversal of actions and shifts of attention underlie higher levels of cognition common to both species. Accepted after revision: 5 May 2001 Electronic Publication