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.     

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.     

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.     

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.     

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.     

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.     

Color preferences in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes)

Color plays an important biological role in the lives of many animals, with some species exhibiting preferences for certain colors over others. This study explored the color preferences of two species of ape, which, like humans, possess trichromatic color vision. Six western lowland gorillas, and six chimpanzees, housed in Belfast Zoological Gardens, were exposed to three stimuli (cloths, boxes, sheets of acetate) in red, blue, and green. Six stimuli of the same nature, in each of the three colors, were provided to both species for 5 days per stimulus. The amount of interest that the animals showed toward each stimulus of each color was recorded for 1 hr. Results showed that the apes, both when analyzed as two separate groups, and when assessed collectively, showed significant color preferences, paying significantly less attention to the red-, than to the blue- or green-colored stimuli. The animals' interest in the blue- and green-colored stimuli did not differ significantly. Overall, the findings suggest that gorillas and chimpanzees, our closest living relatives, may harbor color preferences comparable to those of humans and other species.     

Quantity estimation and comparison in western lowland gorillas (Gorilla gorilla gorilla)

We investigated the quantity judgment abilities of two adult male western lowland gorillas (Gorilla gorilla gorilla) by presenting discrimination tasks on a touch-screen computer. Both gorillas chose the larger quantity of two arrays of dot stimuli. On some trials, the relative number of dots was congruent with the relative total area of the two arrays. On other trials, number of dots was incongruent with area. The gorillas were first tested with static dots, then with dots that moved within the arrays, and finally on a task where they were required to discriminate numerically larger subsets within arrays of moving dots. Both gorillas achieved above-chance performance on both congruent and incongruent trials with all tasks, indicating that they were able to use number as a cue even though ratio of number and area significantly controlled responding, suggesting that number was not the only relevant dimension that the gorillas used. The pattern of performance was similar to that found previously with monkeys and chimpanzees but had not previously been demonstrated in gorillas within a computerized test format, and with these kinds of visual stimuli.     

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.     

Imitative learning by captive western lowland gorillas (Gorilla gorilla gorilla) in a simulated food-processing task

Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.     

Hand preference for food processing in wild western lowland gorillas (Gorilla gorilla gorilla)

Western lowland gorillas (Gorilla gorilla gorilla) have not previously been represented in studies of laterality in wild great apes. The discovery of swampy clearings frequented by gorillas in northern Congo has provided the first opportunity to redress this imbalance. Hand preference data are presented from 33 gorillas in seated and standing postures, covering the procurement and processing of 2 to 4 plant species. Levels of hand preference exhibited were low. When data from all postures and plant species were pooled, 33% of gorillas showed hand preferences in excess of chance. In the standing posture, more gorillas exhibited significant left-hand preferences than right, but an overall population-level bias was not evident.     

Preference for free or forced choice in Sumatran orangutans (Pongo abelii)

Empirical investigations of humans, pigeons, rats, and monkeys have indicated that these species will select free over forced choice, even when faced with identical outcomes. However, the same has yet to be quantitatively confirmed in nonhuman great apes. This experiment is the first systematic investigation of preference for free or forced choice in great apes using a paradigm in which extraneous variables are highly controlled. Three orangutans were given a choice of one of two virtual routes, one that provided a choice and one that did not via a touchscreen computer program. Choice of either route was rewarded with the same type and quantity of food. Initial results indicated a preference for free choice across all three participants. However, in two control conditions, orangutans' preferences varied, suggesting a weaker tendency to exercise choice than species previously tested. We suggest further investigation of preference for free and forced choice in orangutans and other great apes through alternative experimental paradigms that focus on increasing the fidelity of free and forced choice options.     

Matching familiar and unfamiliar faces on internal and external features

Two experiments are reported in which subjects were asked to match a photograph of a complete face and a simultaneously presented photograph of internal or external features of a face, deciding whether or not the two photographs were pictures of the same person. In experiment 1 'same' pairs were derived from different pictures of the same face, so that subjects had to match the faces and not the particular photographs used. Matches based on internal features were found to be faster for familiar than for unfamiliar faces, whereas there was no difference in reaction time between matches based on the external features of familiar and unfamiliar faces. Faster matching of internal features of familiar faces was found to hold equally for pairs of photographs that differed in orientation of the face or in facial expression. In experiment 2 'same' pairs were derived from the same photographs, which gave subjects the choice of matching on the basis of the features of the depicted faces or matching the photographs. Reaction times were faster than in experiment 1, and there were no differences between familiar and unfamiliar faces. The study confirms reports of differential saliency of the internal features of familiar faces, and shows that this only holds when stimuli are treated as faces. The finding thus reflects properties of structural rather than pictorial codes.     

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.     

Mirror self-recognition in gorillas (Gorilla gorilla gorilla): a review and evaluation of mark test replications and variants

Animal Cognition - Mirror self-recognition (MSR), widely regarded as an indicator of self-awareness, has not been demonstrated consistently in gorillas. We aimed to examine this issue by setting...     

Gestural communication of the gorilla (Gorilla gorilla): repertoire, intentionality and possible origins

Social groups of gorillas were observed in three captive facilities and one African field site. Cases of potential gesture use, totalling 9,540, were filtered by strict criteria for intentionality, giving a corpus of 5,250 instances of intentional gesture use. This indicated a repertoire of 102 gesture types. Most repertoire differences between individuals and sites were explicable as a consequence of environmental affordances and sampling effects: overall gesture frequency was a good predictor of universality of occurrence. Only one gesture was idiosyncratic to a single individual, and was given only to humans. Indications of cultural learning were few, though not absent. Six gestures appeared to be traditions within single social groups, but overall concordance in repertoires was almost as high between as within social groups. No support was found for the ontogenetic ritualization hypothesis as the chief means of acquisition of gestures. Many gestures whose form ruled out such an origin, i.e. gestures derived from species-typical displays, were used as intentionally and almost as flexibly as gestures whose form was consistent with learning by ritualization. When using both classes of gesture, gorillas paid specific attention to the attentional state of their audience. Thus, it would be unwarranted to divide ape gestural repertoires into ‘innate, species-typical, inflexible reactions’ and ‘individually learned, intentional, flexible communication’. We conclude that gorilla gestural communication is based on a species-typical repertoire, like those of most other mammalian species but very much larger. Gorilla gestures are not, however, inflexible signals but are employed for intentional communication to specific individuals.     

Single-trial learning of "what" and "who" information in a gorilla (Gorilla gorilla gorilla): implications for episodic memory

Single-trial learning and long-term memory of "what" and "who" information were examined in an adult gorilla (Gorilla gorilla gorilla). We presented the gorilla with a to-be-remembered food item at the time of study. In Experiment 1, following a retention interval of either approximately 7 min or 24 h, the gorilla responded with one of five cards, each corresponding to a particular food. The gorilla was accurate on 70% of the short retention-interval trials and on 82% of the long retention-interval trials. In Experiment 2, the food stimulus was provided by one of two experimenters, each of whom was represented by a card. The gorilla identified the food (55% of the time) and the experimenter (82% of the time) on the short retention-interval trials. On the long retention-interval trials, the gorilla was accurate for the food (73%) and for the person (87%). The results are interpreted in light of theories of episodic memory. Electronic Publication     

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.     

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.