Piagetian conservation of discrete quantities in bonobos (Pan paniscus), chimpanzees (Pan troglodytes), and orangutans (Pongo pygmaeus)

This study investigated whether physical discreteness helps apes to understand the concept of Piagetian conservation (i.e. the invariance of quantities). Subjects were four bonobos, three chimpanzees, and five orangutans. Apes were tested on their ability to conserve discrete/continuous quantities in an over-conservation procedure in which two unequal quantities of edible rewards underwent various transformations in front of subjects. Subjects were examined to determine whether they could track the larger quantity of reward after the transformation. Comparison between the two types of conservation revealed that tests with bonobos supported the discreteness hypothesis. Bonobos, but neither chimpanzees nor orangutans, performed significantly better with discrete quantities than with continuous ones. The results suggest that at least bonobos could benefit from the discreteness of stimuli in their acquisition of conservation skills.     

Inferences about the location of food in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus)

Bonobos (Pan paniscus; n = 4), chimpanzees (Pan troglodytes; n = 12), gorillas (Gorilla gorilla; n = 8), and orangutans (Pongo pygmaeus; n = 6) were presented with 2 cups (1 baited) and given visual or auditory information about their contents. Visual information consisted of letting subjects look inside the cups. Auditory information consisted of shaking the cup so that the baited cup produced a rattling sound. Subjects correctly selected the baited cup both when they saw or heard the food. Nine individuals were above chance in both visual and auditory conditions. More important, subjects as a group selected the baited cup when only the empty cup was either shown or shaken, which means that subjects chose correctly without having seen or heard the food (i.e., inference by exclusion). Control tests showed that subjects were not more attracted to noisy cups, avoided shaken noiseless cups, or learned to use auditory information as a cue during the study. It is concluded that subjects understood that the food caused the noise, not simply that the noise was associated with the food.     

Discrete quantity judgments in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, Pongo pygmaeus): the effect of presenting whole sets versus item-by-item

The authors examined quantity-based judgments for up to 10 items for simultaneous and sequential whole sets as well as for sequentially dropped items in chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), bonobos (Pan paniscus), and orangutans (Pongo pygmaeus). In Experiment 1, subjects had to choose the larger of 2 quantities presented in 2 separate dishes either simultaneously or 1 dish after the other. Representatives of all species were capable of selecting the larger of 2 quantities in both conditions, even when the quantities were large and the numerical distance between them was small. In Experiment 2, subjects had to select between the same food quantities sequentially dropped into 2 opaque cups so that none of the quantities were ever viewed as a whole. The authors found some evidence (albeit weaker) that subjects were able to select the larger quantity of items. Furthermore, the authors found no performance breakdown with the inclusion of certain quantities. Instead, the ratio between quantities was the best performance predictor. The authors conclude that quantity-based judgments rely on an analogical system, not a discrete object file model or perceptual estimation mechanism, such as subitizing.     

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.     

Tracking the displacement of objects: a series of tasks with great apes (Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus) and young children (Homo sapiens)

The authors administered a series of object displacement tasks to 24 great apes and 24 30-month-old children (Homo sapiens). Objects were placed under 1 or 2 of 3 cups by visible or invisible displacements. The series included 6 tasks: delayed response, inhibition test, A not B, rotations, transpositions, and object permanence. Apes and children solved most tasks performing at comparable levels except in the transposition task, in which apes performed better than children. Ape species performed at comparable levels in all tasks except in single transpositions, in which chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) performed better than gorillas (Gorilla gorilla) and orangutans (Pongo pygmeaus). All species found nonadjacent trials and rotations especially difficult. The number of elements that changed locations, the type of displacement, and having to inhibit predominant reaching responses were factors that negatively affected the subjects' performance.     

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.     

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.     

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.     

Spontaneous spatial constructions by chimpanzees (Pan troglodytes,Pan paniscus)

We investigated chimpanzees’ spontaneous spatial constructions with objects. Two were common chimpanzees ages 6 and 18 years and one was an 11‐year‐old bonobo raised in a very enriched human environment from very early in life. Chimpanzees’ products and procedures were quite advanced when constructing functional spatial relations by placing objects in or on each other, but not when constructing nonfunctional spatial relations by placing objects next to each other. This difference was found in all their constructions, including their spatial correspondences and spatial symmetries. Compared to those of younger nonencultured chimpanzees all their spatial constructions were more advanced. Compared to human infants, their functional spatial constructions were also more advanced but their nonfunctional spatial constructions were less advanced.     

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.

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.     

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.     

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.     

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.     

Spontaneous logicomathematical constructions by chimpanzees (Pan troglodytes, P. paniscus)

Two experiments investigated the spontaneous construction of precursory logicomathematical operations by human-enculturated and language-reared chimpanzees (Pan troglodytes, Pan paniscus) when they were interacting freely with objects. In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with sets of six objects. Chimpanzees constructed equivalence, order and reversibility relations within single sets of objects as well as between two or three contemporaneous sets of objects. The chimpanzees’ logicomathematical operations were more advanced, including infrequent and minimal operations on three sets, than those of some previously investigated younger nonenculturated common chimpanzees. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with sets of 12 objects. Chimpanzees constructed more advanced operations on single sets, but not on contemporaneous sets. The results suggest partial convergence and partial divergence between development of logicomathematical cognition in chimpanzees and humans. Received: 20 January 1999 / Accepted after revision: 9 August 1999     

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.     

Relative numerousness judgment and summation in young, middle-aged, and older adult orangutans (Pongo pygmaeus abelii and Pongo pygmaeus pygmaeus)

The ability to select the larger of two quantities ranging from 1 to 5 (relative numerousness judgment [RNJ[) and the ability to select the larger of two pairs of quantities with each pair ranging from 1 to 8 (summation) were evaluated in young, middle-aged, and older adult orangutans (7 Pongo pygmaeus abelii and 2 Pongo pygmaeus pygmaeus). Summation accuracy and RNJ were similar to those of previous reports in apes; however, the pattern of age-related differences with regard to these tasks was different from that previously reported in gorillas. Older orangutans were less accurate than the young and middle-aged for RNJ, and summation accuracy was equivalent among age groups. Evidence was found to suggest that the young and middle-aged based their selection of the largest quantity pair on both quantities within each pair during the summation task. These results show a relationship between subject age and the quantitative abilities of adult orangutans.     

Spontaneous classification in action by a human-enculturated and language-reared bonobo (Pan paniscus) and common chimpanzees (Pan troglodytes).

The authors investigated the formal features of spontaneous manipulations used by 1 bonobo (Pan paniscus) and 2 common chimpanzees (Pan troglodytes) to classify objects in action. Chimpanzees' manipulations were evenly split between serial, one-at-a-time acts on 1 object and parallel, two-at-a-time acts on 2 or 3 objects. Chimpanzees systematically combined their manipulations into routines to generate class-consistent categories of objects. Their routines featured much reproduction of the same manipulations, planful acts that anticipated follow-up manipulations, and manipulations that were reciprocal to each other to accomplish an end. In all these respects, chimpanzees' manipulations were similar to those of 2-year-old human infants. In others they differed. Chimpanzees' routines were mainly based on a linear integration of manipulations. Classifying in action was only infrequently produced by hierarchically integrated routines.     

Physical intuitions about support relations in monkeys (Macaca fuscata) and apes (Pan troglodytes)

Nonhuman primates, like humans, have demonstrated various physical intuitions. Cacchione and Krist (2004) examined chimpanzees' intuitions about support relations with the violation-of-expectation task. They reported that the chimpanzees possessed intuitions about support, but their intuitions differed from those of humans in part; they were sensitive to "contact/no-contact" and "amount of contact" but not "type of contact" rule. To further explore intuitions about support in nonhuman primates, we conducted similar experiments on monkeys (Japanese monkeys) and apes (chimpanzees). In three experiments, we presented physically possible and impossible events of different support relations to the participants and measured their looking times. The results reveal that the chimpanzees and monkeys detect the violations of "contact/no-contact" and "amount of contact" but not "type of contact" variable. Therefore, the apes and monkeys possess similar intuitions; however, these intuitions differ in part from those of humans. The present study provides new and corroborative evidence of intuitions about support in nonhuman primates. However, this again leads to the question of distinctive understanding about support relations among primate species.