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.     

The effect of response contingencies on scale model task performance by chimpanzees (Pan troglodytes)

The effects of modified procedures on chimpanzees' (Pan troglodytes) performance in a scale model comprehension task were examined. Seven chimpanzees that previously participated in a task in which they searched an enclosure for a hidden item after watching an experimenter hide a miniature item in the analogous location in a scale model were retested under procedures incorporating response costs. In Experiment 1, chimpanzees were trained under procedures that rewarded only item retrievals occurring on the 1st search attempt. During test trials, 6 chimpanzees performed above chance, including 4 that were previously unsuccessful under the original procedures (V. A. Kuhlmeier, S. T. Boysen, & K. L. Mukobi, 1999). Experiment 2 compared performance under the new and original procedures. Results indicated that for some chimpanzees, performance depended on procedures that decreased the use of competing search strategies and encouraged strategies based on information from the scale model.     

What meaning means for same and different: Analogical reasoning in humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta)

Thus far, language- and token-trained apes (e.g., D. Premack, 1976; R. K. R. Thompson, D. L. Oden, & S. T. Boysen, 1997) have provided the best evidence that nonhuman animals can solve, complete, and construct analogies, thus implicating symbolic representation as the mechanism enabling the phenomenon. In this study, the authors examined the role of stimulus meaning in the analogical reasoning abilities of three different primate species. Humans (Homo sapiens), chimpanzees (Pan troglodytes), and rhesus monkeys (Macaca mulatta) completed the same relational matching-to-sample (RMTS) tasks with both meaningful and nonmeaningful stimuli. This discrimination of relations-between-relations serves as the basis for analogical reasoning. Meaningfulness facilitated the acquisition of analogical matching for human participants, whereas individual differences among the chimpanzees suggest that meaning can either enable or hinder their ability to complete analogies. Rhesus monkeys did not succeed in the RMTS task regardless of stimulus meaning, suggesting that their ability to reason analogically, if present at all, may be dependent on a dimension other than the representational value of stimuli.     

Mangabeys (<Emphasis Type="Italic">Cercocebus torquatus lunulatus</Emphasis>) solve the reverse contingency task without a modified procedure

Problem solving often relies on generating new responses while inhibiting others, particularly prepotent ones. A paradigm to study inhibitory abilities is the reverse contingency task (Boysen and Berntson in J Exp Psychol Anim Behav Process 21:82–86, 1995), in which two different quantities of food are offered to an individual who receives the array he did not choose. Therefore, mastery of the task demands selecting the smaller quantity to obtain the larger one. Several non-human primates have been tested in the reverse contingency task. To date, only great apes and rhesus monkeys (Macaca mulatta) have succeeded in the original task, with no need of procedural modifications as the large-or-none contingency, correction trials or symbolic stimuli substituting for actual food quantities. Here, four mangabeys were presented with two stimulus arrays of one and four raisins in the context of the reverse contingency task. Three of them learned to perform the task well above chance without a modified procedure. They also reached above-chance performance when presented with two stimulus arrays of zero and four raisins, despite the initial difficulty of choosing a null quantity. After a period of 7–10 months, in which the animals were not tested on any task, all three subjects continued to perform well, even when presented with novel quantity pairs.     

Chimpanzee (Pan troglodytes) relational matching: playing by their own (analogical) rules

Chimpanzees (Pan troglodytes) have been known to exhibit rudimentary abilities in analogical reasoning (Flemming, Beran, Thompson, Kleider, & Washburn, 2008; Gillian, Premack, & Woodruff, 1981; Haun & Call, 2009; Thompson & Oden, 2000; Thompson, Oden, & Boysen, 1997). With a wide array of individual differences, little can be concluded about the species' capacity for analogies, much less their strategies employed for solving such problems. In this study, we examined analogical strategies in 3 chimpanzees using a 3-dimensional search task (e.g., Kennedy & Fragaszy, 2008). Food items were hidden under 1 of 2 or 3 plastic cups of varying sizes. Subsequently, chimpanzees searched for food under the cup of the same relative size in their own set of cups--reasoning by analogy. Two chimpanzees initially appeared to fail the first relational phase of the task. Meta-analyses revealed, however, that they were instead using a secondary strategy not rewarded by the contingencies of the task--choosing on the basis of the same relative position in the sample. Although this was not the intended strategy of the task, it was nonetheless analogical. In subsequent phases of the task, chimpanzees eventually learned to shift their analogical reasoning strategy to match the reward contingencies of the task and successfully choose on the basis of relative size. This evidence not only provides support for the analogical ape hypothesis (Thompson & Oden, 2000), but also exemplifies how foundational conceptually mediated analogical behavior may be for the chimpanzee.     

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.     

Pointing At Smaller Food Amounts In An Analogue Of Boysen And Berntson's (1995) Procedure

Boysen and Berntson (1995) showed that apes could not learn to point to a small amount of candy in order to get a larger amount when pointing to the larger amount was reinforced by receipt of the alternate, smaller amount. They explained this result as an unlearned predisposition to reach for higher value foods that overrides the effects of reinforcement. This report tests their thesis. In the first condition, 3 monkeys chose between one raisin held in one hand by the experimenter and four raisins held in the other hand. If a monkey pointed at four raisins, it received one. If it pointed at one, it received four. Over ten 20-trial sessions, no monkey learned to point at the one-raisin alternative, a result similar to that of Boysen and Berntson. In the second condition, pointing at one raisin still produced four; however, pointing at four raisins now produced no reinforcement. In five 20-trial sessions, all monkeys learned to point at one raisin in order to get four. This finding demonstrates that at least in monkeys there is no predisposition to reach for higher value foods that cannot be readily overridden by reinforcement contingencies, and casts doubt on Boysen and Berntson's claim to have demonstrated such a process in apes.     

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).     

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.     

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.     

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.     

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.     

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.     

Inferences by exclusion in the great apes: the effect of age and species

This study investigated the ability of chimpanzees, gorillas, orangutans, and bonobos to make inferences by exclusion using the procedure pioneered by Premack and Premack (Cognition 50:347–362, 1994) with chimpanzees. Thirty apes were presented with two different food items (banana vs. grape) on a platform and covered with identical containers. One of the items was removed from the container and placed between the two containers so that subjects could see it. After discarding this item, subjects could select between the two containers. In Experiment 1, apes preferentially selected the container that held the item that the experimenter had not discarded, especially if subjects saw the experimenter remove the item from the container (but without seeing the container empty). Experiment 3 in which the food was removed from one of the containers behind a barrier confirmed these results. In contrast, subjects performed at chance levels when a stimulus (colored plastic chip: Exp. 1; food item: Exp. 2 and Exp. 3) designated the item that had been removed. These results indicated that apes made inferences, not just learned to use a discriminative cue to avoid the empty container. Apes perceived and treated the item discarded by the experimenter as if it were the very one that had been hidden under the container. Results suggested a positive relationship between age and inferential ability independent of memory ability but no species differences.This contribution is part of the special issue “Animal Logics” (Watanabe and Huber 2006).     

How great apes perform on a modified trap-tube task

To date, neither primates nor birds have shown clear evidence of causal knowledge when attempting to solve the trap tube task. One factor that may have contributed to mask the knowledge that subjects may have about the task is that subjects were only allowed to push the reward away from them, which is a particularly difficult action for primates in certain problem solving situations. We presented five orangutans (Pongo pygmaeus), two chimpanzees (Pan troglodytes), two bonobos (Pan paniscus), and one gorilla (Gorilla gorilla) with a modified trap tube that allowed subjects to push or rake the reward with the tool. In two additional follow-up tests, we inverted the tube 180° rendering the trap nonfunctional and also presented subjects with the original task in which they were required to push the reward out of the tube. Results showed that all but one of the subjects preferred to rake the reward. Two orangutans and one chimpanzee (all of whom preferred to rake the reward), consistently avoided the trap only when it was functional but failed the original task. These findings suggest that some great apes may have some causal knowledge about the trap-tube task. Their success, however, depended on whether they were allowed to choose certain tool-using actions. Electronic Supplementary Material Supplementary material is available for this article at     

Prospective memory in children and chimpanzees

Prospective memory (PM) involves remembering to do something at a specific time in the future. Here, we investigate the beginnings of this ability in young children (3-year-olds; Homo sapiens) and chimpanzees (Pan troglodytes) using an analogous task. Subjects were given a choice between two toys (children) or two food items (chimpanzees). The selected item was delivered immediately, whereas the unselected item was hidden in an opaque container. After completing an ongoing quantity discrimination task, subjects could request the hidden item by asking for it (children) or by pointing to the container and identifying the item on a symbol board (chimpanzees). Children and chimpanzees showed evidence of prospective-like memory in this task, as evidenced by successful retrieval of the item at the end of the task, sometimes spontaneously with no prompting from the experimenter. These findings contribute to our understanding of PM from an ontogenetic and comparative perspective.     

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.     

Less is more: executive function and symbolic representation in preschool children

Executive function is recognized as a critical component of children's cognitive and social development. In two studies, a measure of executive function that had been used in research with chimpanzees was adapted for preschoolers. On this task, called Less Is More, children must point to a smaller reward (two candies) to receive a larger reward (five candies). In Study 1 (N= 101), performance was significantly related to age (3 vs. 4), verbal ability, and established measures of executive function. In Study 2 (N= 128), symbolic representations substituted for real candies in this task. Three-year-olds' performance improved significantly as a function of symbolic distancing. This research has implications for the role of symbol systems in the development of executive control over thought and action.     

Perceptual learning on inspection time and motion perception

Perceptual learning on simple perceptual tasks is interpreted as plasticity of neuronal populations in the sensory cortex (M. Fahle & T. Poggio, 2002). The authors examined individual differences on perceptual learning for 2 tasks-inspection time (IT) and a motion direction discrimination task that was instantiated as random dot kinematograms. The authors' main questions were whether individual differences in perceptual learning were consistent across the 2 tasks and whether perceptual learning correlated with cognitive abilities test scores. In all, 56 young adults completed 16 threshold estimations on 1 of 2 orthogonal versions of each task. Then, the authors made 2 further threshold estimations for the untrained, orthogonal version. Participants also completed a battery of 6 cognitive abilities tests that measured fluid ability (Gf) and perceptual speed (Gs). Perceptual learning was demonstrated for both tasks, but the degree of learning across tasks was not characteristic of the individual. Learning on IT correlated with Gs (r = .35), but learning on the motion direction discrimination task was unrelated to cognitive ability. Correlations of IT with cognitive measures were stable over the training period. IT was correlated with both the motion direction discrimination task (r = -.39) and an unmasked line length judgment task (r = -.31). The authors concluded that perceptual learning on IT correlates with cognitive abilities test scores, that correlations of IT with cognitive abilities test scores are stable as task performance improves with practice, and that the IT task is psychologically complex.