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.     

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.     

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.     

Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

Erratum to: Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

Spontaneous use of tools as straws in great apes

Great apes can use multiple tools to extract food embedded in substrates and can invent new ways to exploit those resources. We tested five bonobos, five chimpanzees, and six orangutans in a task in which they had to use (and modify) a tool as a straw to drink the juice located inside a container. Experiment 1 showed that four orangutans and one chimpanzee invented the use of a piece of electric cable to get the juice. Experiment 2 investigated whether subjects could transform a non-functional hose into a functional one by removing blockages that impeded the free flow of juice. Orangutans outperformed chimpanzees and bonobos by differentially removing those blockages that prevented the flow of juice, often doing so before attempting to extract the juice. In Experiment 3, we presented chimpanzees and orangutans with four 3-tool sets (each tool set contained a single straw-like tool) and allowed them to select one tool. Unlike chimpanzees, orangutans succeeded in selecting the straw-like tool above chance levels without having to physically manipulate it. We suggest that orangutans’ superior performance is related to their greater reliance on mouth actions during foraging. Experiment 4 investigated whether orangutans were also capable of selecting the suitable tool not by its appearance, but by the effects that it produced. After witnessing the experimenter blow bubbles or absorb liquid with a functional tool but fail to accomplish the same thing with the non-functional tool, orangutans failed to select the functional tool above chance levels.     

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.     

Reaching around barriers: the performance of the great apes and 3–5-year-old children

Inhibitory control has been suggested as a key predictive measure of problem-solving skills in human and nonhuman animals. However, there has yet to be a direct comparison of the inhibitory skills of the nonhuman apes and their development in human children. We compared the inhibitory skills of all great ape species, including 3–5-year-old children in a detour-reaching task, which required subjects to avoid reaching directly for food and instead use an indirect reaching method to successfully obtain the food. We tested 22 chimpanzees, 18 bonobos, 18 orangutans, 6 gorillas and 42 children. Our sample included chimpanzees, bonobos and orangutans housed in zoos (N = 27) and others housed in sanctuaries in their native habitats (N = 37). Overall, orangutans were the most skilful apes, including human children. As expected older children outperformed younger children. Sanctuary chimpanzees and bonobos outperformed their zoo counterparts whereas there was no difference between the two orangutan samples. Most zoo chimpanzees and bonobos failed to solve the original task, but improved their performance with additional training, although the training method determined to a considerable extent the level of success that the apes achieved in a transfer phase. In general, the performance of the older children was far from perfect and comparable to some of the nonhuman apes tested.     

What does an intermediate success rate mean? An analysis of a Piagetian liquid conservation task in the great apes

The study investigates what an intermediate success rate means in bonobos, chimpanzees, and orangutans. Apes participated in liquid conservation experiments where they had to track the larger of two different quantities of juice after various kinds of transformations [Suda, C., & Call, J. (2004). Piagetian liquid conservation in the great apes (Pan paniscus, Pan troglodytes, and Pongo pygmaeus). Journal of Comparative Psychology, 118, 265-279). When making a decision, apes sometimes demonstrated hesitant behavior, concurrently pointing to both alternatives or successively changing their choice. Moderately successful apes showed more hesitation than highly successful or unsuccessful apes. The results are consistent with the cognitive conflict model: The experiments created a higher degree of cognitive conflict on moderately successful apes than on very successful or unsuccessful apes. This indicates that an intermediate performance reflects the joint operation and potential conflict between two different cognitive strategies (identity and appearance) inherent to the Piagetian conservation task.     

Summation in the chimpanzee (Pan troglodytes)

In this research, we asked whether 2 chimpanzee (Pan troglodytes) subjects could reliably sum across pairs of quantities to select the greater total. Subjects were allowed to choose between two trays of chocolates. Each tray contained two food wells. To select the tray containing the greater number of chocolates, it was necessary to sum the contents of the food wells on each tray. In experiments where food wells contained from zero to four chocolates, the chimpanzees chose the greater value of the summed wells on more than 90% of the trials. In the final experiment, the maximum number of chocolates assigned to a food well was increased to five. Choice of the tray containing the greater sum still remained above 90%. In all experiments, subjects reliably chose the greater sum, even though on many trials a food well on the "incorrect" tray held more chocolates than either single well on the "correct" tray. It was concluded that without any known ability to count, these chimpanzees used some process of summation to combine spatially separated quantities. Speculation regarding the basis for summation includes consideration of perceptual fusion of pairs of quantities and subitization.     

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.     

Great apes' understanding of other individuals' line of sight

Previous research has shown that many social animals follow the gaze of other individuals. However, knowledge about how this skill differs between species and whether it shows a relationship with genetic distance from humans is still fragmentary. In the present study of gaze following in great apes, we manipulated the nature of a visual obstruction and the presence/absence of a target. We found that bonobos, chimpanzees, and gorillas followed gaze significantly more often when the obstruction had a window than when it did not, just as human infants do. Additionally, bonobos and chimpanzees looked at the experimenter's side of a windowless obstruction more often than the other species. Moreover, bonobos produced more double looks when the barrier was opaque than when it had a window, indicating an understanding of what other individuals see. The most distant human relatives studied, orangutans, showed few signs of understanding what another individual saw. Instead, they were attracted to the target's location by the target's presence, but not by the experimenter's gaze. Great apes' perspective-taking skills seem to have increased in the evolutionary lineage leading to bonobos, chimpanzees, and humans.     

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.     

A comparison of temperament in nonhuman apes and human infants

The adaptive behavior of primates, including humans, is often mediated by temperament. Human behavior likely differs from that of other primates in part due to temperament. In the current study we compared the reaction of bonobos, chimpanzees, orangutans, and 2.5-year-old human infants to novel objects and people - as a measure of their shyness-boldness, a key temperamental trait. Human children at the age of 2.5 years avoided novelty of all kinds far more than the other ape species. This response was most similar to that seen in bonobos and least like that of chimpanzees and orangutans. This comparison represents a first step in characterizing the temperamental profiles of species in the hominoid clade, and these findings are consistent with the hypothesis that human temperament has evolved since our lineage diverged from the other apes in ways that likely have broad effects on behavior. These findings also provide new insights into how species differences in ecology may shape differences in temperament.     

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.     

Great apes’ performance in discriminating weight and achromatic color

Much work has been done on visual discrimination in primates over the past decade. In contrast, very little is known about the relevance of non-visual information in discrimination learning. We investigated weight and achromatic color (color, henceforth) discrimination in bonobos, gorillas and orangutans, using the exchange paradigm in which subjects have to give objects to the experimenter in order to receive a reward. Unlike previous studies, subjects were not trained to lift objects because lifting the objects was an integral part of the exchange procedure. This methodology also allowed us a direct comparison between visual and weight discrimination. We presented 12 subjects (5 bonobos, 2 gorillas and 5 orangutans) with two sets of objects corresponding to two conditions. The objects in the color condition (white/black) differed only in color and those in the weight condition (light/heavy) differed only in weight. Five apes learned to discriminate weight and six to discriminate color. Subjects learned color discrimination faster than weight discrimination. Our results suggest that bonobos and orangutans are sensitive to differences in weight and able to learn discriminating objects that differ in this property.     

Behavioral cues that great apes use to forage for hidden food

We conducted three studies to examine whether the four great ape species (chimpanzees, bonobos, gorillas, and orangutans) are able to use behavioral experimenter-given cues in an object-choice task. In the subsequent experimental conditions subjects were presented with two eggs, one of which contained food and the other did not. In Study 1 the experimenter examined both eggs by smelling or shaking them, but only made a failed attempt to open (via biting) the egg containing food. In a control condition, the experimenter examined and attempted to open both eggs, but in reverse order to control for stimulus enhancement. The apes significantly preferred the egg that was first examined and then bitten, but had no preference in a baseline condition in which there were no cues. In Study 2, we investigated whether the apes could extend this ability to cues not observed in apes so far (i.e., attempting to pull apart the egg), as well as whether they made this discrimination based on the function of the action the experimenter performed. Subjects significantly preferred eggs presented with this novel cue, but did not prefer eggs presented with a novel but functionally irrelevant action. In Study 3, apes did not interpret human actions as cues to food-location when they already knew that the eggs were empty. Thus, great apes were able to use a variety of experimenter-given cues associated with foraging actions to locate hidden food and thereby were partially sensitive to the general purpose underlying these actions.     

Effects of element separation on perceptual grouping by humans (Homo sapiens) and chimpanzees (Pan troglodytes): perception of Kanizsa illusory figures

The processing of Kanizsa-square illusory figures was studied in two experiments with four humans and two chimpanzees. Subjects of the two species were initially trained to select a Kanizsa-square illusory figure presented in a computerized two-alternative forced choice task. After training, adding narrow closing segments to the pacman inducers that composed the Kanisza illusory figures lowered performance in both chimpanzees and humans, suggesting that the discrimination could be controlled by the perception of illusory forms. A second experiment assessed transfer of performance with five sets of figures in which the size of the inducers and their separation were manipulated. Only for chimpanzees was performance directly controlled by separation, suggesting that chimpanzees are more sensitive than humans to the separation between visual elements. Accepted after revision: 15 August 2001 Electronic Publication     

An obedient orangutan (Pongo abelii) performs perfectly in peripheral object-choice tasks but fails the standard centrally presented versions

Mulcahy and Call (2009) found that bonobos (Pan paniscus) and chimpanzees (Pan troglodytes) but not orangutans (Pongo pygmaeus) perform significantly better in a peripheral version of the object-choice task compared to the original central version. Orangutans may have failed because they avoided direct eye contact with the experimenter when the cue was given. We investigated this possibility by conducting peripheral and central object choice tasks with an obedient orangutan (Pongo abelii) whom the experimenter could elicit eye contact with in each trial. In contrast to Mulcahy and Call's findings, the subject only failed the object choice task when tested with the central and not the peripheral version. We investigated whether success was because of the greater distance the subject was required to move in order to make a choice in peripheral trials. Results show that this was an unlikely factor in the subject's success. We discuss our findings in relation to previous and future object-choice research.     

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.