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.     

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.     

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.     

Orangutans (Pongo abelii) "play the odds": information-seeking strategies in relation to cost, risk, and benefit

Recent research has examined whether animals possess metacognition, or the ability to monitor their knowledge states. However, the extent to which animals actively control their knowledge states is still not well delineated. Although organisms might be capable of seeking information when it is lacking, it does not mean that it is always adaptive to do so. In the present set of experiments, we examined the flexibility of this behavior in captive orangutans (Pongo abelii; two adults and one juvenile) in a foraging task, by varying the necessity of information-seeking, the cost associated with it, the likelihood of error, and the value of the reward. In Experiment 1, subjects searched for information most often when it was "cheapest" energetically. In Experiment 2, subjects searched for information most often when the odds of making an error were the greatest. In Experiment 3, subjects searched for information more when the reward was doubled in value. In Experiment 4, adult subjects adapted to risk/benefit trade-offs in their searching behavior. In every experiment, subjects sought information more often when they needed it than when they already knew the solution to the problem. Therefore, the current research suggests that information-seeking behavior in orangutans shows a sophisticated level of flexibility, comparable to that seen in human children, as they appear to "play the odds" when making the decision to seek information or not.     

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.     

Levels of abstraction in orangutan (Pongo abelii) categorization

Levels of abstraction have rarely been manipulated in studies of natural concept formation in nonhumans. Isolated examples have indicated that animals, relative to humans, may learn concepts at varying levels of abstraction with differential ease. The ability of 6 orangutans (Pongo abelii) of various ages to make natural concept discriminations at 3 levels of abstraction was therefore investigated. The orangutans were rewarded for selecting photos of orangutans instead of humans and other primates (concrete level), primates instead of other animals (intermediate level), and animals instead of nonanimals (abstract level) in a 2-choice touch screen procedure. The results suggest that, like a gorilla (Gorilla gorilla gorilla) tested previously (Vonk & MacDonald, 2002), orangutans can learn concepts at each level of abstraction, and unlike other nonhumans, most of these subjects rapidly learned the intermediate level discrimination.     

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.     

Orangutans (Pongo spp.) may prefer tools with rigid properties to flimsy tools

Preference for tools with either rigid or flexible properties was explored in orangutans (Pongo spp.) through an extension of D. J. Povinelli, J. E. Reaux, and L. A. Theall's (2000) flimsy-tool problem. Three captive orangutans were presented with three unfamiliar pairs of tools to solve a novel problem. Although each orangutan has spontaneously used tools in the past, the tools presented in this study were novel to the apes. Each pair of tools contained one tool with rigid properties (functional) and one tool with flimsy properties (nonfunctional). Solving the problem required selection of a rigid tool to retrieve a food reward. The functional tool was selected in nearly all trials. Moreover, two of the orangutans demonstrated this within the first test trials with each of the three tool types. Although further research is required to test this statistically, it suggests either a preexisting preference for rigid tools or comprehension of the relevant features required in a tool to solve the task. The results of this study demonstrate that orangutans can recognize, or learn to recognize, relevant tool properties and can choose an appropriate tool to solve a problem.     

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.     

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.     

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.     

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.     

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.     

Inferential reasoning by exclusion in great apes, lesser apes, and spider monkeys

Using the cups task, in which subjects are presented with limited visual or auditory information that can be used to deduce the location of a hidden reward, Call (2004) found prima facie evidence of inferential reasoning by exclusion in several great ape species. One bonobo (Pan paniscus) and two gorillas (Gorilla gorilla) appeared to make such inferences in both the visual and auditory domains. However, common chimpanzees (Pan troglodytes) were successful only in the visual domain, and Bornean orangutans (Pongo pygmaeus) in neither. The present research built on this paradigm, and Experiment 1 yielded prima facie evidence of inference by exclusion in both domains for two common chimpanzees, and in the visual domain for two Sumatran orangutans (Pongo abelii). Experiments 2 and 3 demonstrated that two specific associative learning explanations could not readily account for these results. Because an important focus of the program of research was to assess the cognitive capacities of lesser apes (family Hylobatidae), we modified Call's original procedures to better suit their attentional and dispositional characteristics. In Experiment 1, testing was also attempted with three gibbon genera (Symphalangus, Nomascus, Hylobates), but none of the subjects completed the standard task. Further testing of three siamangs (Symphalangus syndactylus) and a spider monkey (Ateles geoffroyi) using a faster method yielded prima facie evidence of inferential reasoning by exclusion in the visual domain among the siamangs (Experiment 4).     

Quantity matching by an orangutan (Pongo abelii)

An adult male orangutan (Pongo abelii) was presented with a series of delayed matching-to-sample (DMTS) tasks in which he was to match images based on (a) the number of individual animals depicted in the photograph (from 1 to 4), (b) the number of abstract shapes presented in the stimulus (from 1 to 4), or (c) the number of dots presented in the stimulus (from 1 to 4, 4–7, or 7–10). The spatial arrangement of the dots and the background color of the stimuli varied, and the size of the dots was manipulated to control for overall ratio of foreground to background. The subject’s performance was not affected by these perceptual features, but was affected by the absolute difference and ratio between number of elements in the comparison stimuli. However, the relationship between these variables and his performance was not always linear as predicted by the analog magnitude model. In addition, the subject showed a high degree of transfer to novel numerosities up to ten, indicating that orangutans are capable of estimating quantity for a greater number of items than can presumably be subtilized by humans.     

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.     

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.     

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.