Estimating and operating on discrete quantities in orangutans (Pongo pygmaeus)

This study investigated the ability of 3 male orangutans (Pongo pygmaeus; 1 subadult, 2 adults) to estimate, compare, and operate on 2 sets of small quantities (1-6 cereal bits). Experiment 1 investigated the orangutans' ability to choose the larger of 2 quantities when they were presented successively as opposed to simultaneously, thus being perceptually unavailable at the time of choice. Experiment 2 investigated the orangutans' ability to select the larger quantity after the original quantities were augmented or reduced. Orangutans were capable of selecting the larger of 2 quantities in Experiment 1. There was also some evidence from Experiment 2, albeit weaker, that orangutans may mentally combine quantities (but not dissociate) to obtain the larger of 2 quantities. This study suggests that orangutans use a representational mechanism (especially when comparing quantities) to select the larger of 2 sets of items.     

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.     

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.     

Visual discrimination in an orangutan (Pongo pygmaeus): measuring visual preference

Although previous studies have confirmed that trained orangutans visually discriminate between mammals and artificial objects, whether orangutans without operant conditioning can discriminate remains unknown. The visual discrimination ability in an orangutan (Pongo pygmaeus) with no experience in operant learning was examined using measures of visual preference. Sixteen color photographs of inanimate objects and of mammals with four legs were randomly presented to an orangutan. The results showed that the mean looking time at photographs of mammals with four legs was longer than that for inanimate objects, suggesting that the orangutan discriminated mammals with four legs from inanimate objects. The results implied that orangutans who have not experienced operant conditioning may possess the ability to discriminate visually.     

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.     

Relative numerousness judgment and summation in young and old Western lowland gorillas

The relationship between age, relative numerousness judgment, and summation was investigated in 11 Western lowland gorillas (Gorilla gorilla gorilla). Experiments 1 and 2 evaluated the gorillas' ability to select the larger of 2 food quantities before and with training. The majority of gorillas did not reliably select the larger quantity in Experiment 1 until receiving training to do so in Experiment 2. Experiment 3 evaluated their ability to select the larger of 2 pairs of quantities. All gorillas selected the larger pair more often than chance, and the old were less accurate and slower than were the young. For most gorillas, performance in relative numerousness judgment with training and summation was comparable with previous reports in chimpanzees and orangutans.     

Social learning by orangutans (Pongo abelii and Pongo pygmaeus) in a simulated food-processing task

Increasing evidence for behavioral differences between populations of primates has created a resurgence of interest in examining mechanisms of information transfer between individuals. The authors examined the social transmission of information in 15 captive orangutans (Pongo abelii and Pongo pygmaeus) using a simulated food-processing task. Experimental subjects were shown 1 of 2 methods for removing a suite of defenses on an "artificial fruit." Control subjects were given no prior exposure before interacting with the fruit. Observing a model provided a functional advantage in the task, as significantly more experimental than control subjects opened the fruit. Within the experimental groups, the authors found a trend toward differences in the actual behaviors used to remove 1 of the defenses. Results support observations from the wild implying horizontal transfer of information in orangutans and show that a number of social learning processes are likely to be involved in the transfer of knowledge in this species.     

An assessment of generalization of imitation in two enculturated orangutans (Pongo pygmaeus)

Deferred imitation of object-related actions and generalization of imitation to similar but not identical tasks were assessed in two human-reared (enculturated) orangutans (Pongo pygmaeus). Each ape displayed low levels of deferred imitation but did not provide evidence of generalization of imitation. Results suggest that enculturated orangutans may not possess social-cognitive abilities similar to that which enculturated chimpanzees are assumed to possess.     

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.     

Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task

Two important elements in problem solving are the abilities to encode relevant task features and to combine multiple actions to achieve the goal. The authors investigated these 2 elements in a task in which gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) had to use a tool to retrieve an out-of-reach reward. Subjects were able to select tools of an appropriate length to reach the reward even when the position of the reward and tools were not simultaneously visible. When presented with tools that were too short to retrieve the reward, subjects were more likely to refuse to use them than when tools were the appropriate length. Subjects were proficient at using tools in sequence to retrieve the reward.     

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.     

Nonenculturated orangutans' (Pongo pygmaeus) use of experimenter-given manual and facial cues in an object-choice task

Several experiments have been performed, to examine whether nonhuman primates are able to make use of experimenter-given manual and facial (visual) cues to direct their attention to a baited object. Contrary to the performance of prosimians and monkeys, great apes repeatedly have shown task efficiency in experiments such as these. However, many great ape subjects used have been "enculturated" individuals. In the present study, 3 nonenculturated orangutans (Pongo pygmaeus) were tested for their ability to use experimenter-given pointing, gazing, and glancing cues in an object-choice task. All subjects readily made use of the pointing gesture. However, when subjects were left with only gazing or glancing cues, their performance deteriorated markedly, and they were not able to complete the task.     

Response strategies in list learning by orangutans (Pongo pygmaeus x P. abelii)

Rhesus monkeys (Macaca mulatta) develop strategies to acquire and execute serial lists (K. B. Swartz & S. A. Himmanen, 2001). Serial probe recognition studies of list memory have demonstrated similarities across monkeys and humans (S. F. Sands & A. A. Wright, 1980). The present study extended the investigation of list learning and memory to determine whether orangutans (Pongo pygmaeus x P. abelii) would show evidence of subjective organization of photographic lists in a manner similar to that shown by humans learning a list of unrelated words (E. Tulving, 1962). No evidence for the effective use of a subjective organization strategy was found, but the orangutans developed a right-to-left spatial response strategy, which emerged during the acquisition of 5-item lists. This strategy was an effective way to reduce the load on working memory when presented with a complex array of items.     

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.     

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.     

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.     

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.     

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.