Addendum to "Summation in the chimpanzee (Pan troglodytes)"

Rumbaugh, Savage-Rumbaugh, and Hegel (1987) reported that two chimpanzees (Pan troglodytes) could select, with better than 90% accuracy, the greater of two paired quantities of chocolate chips. In that study, no one quantity of chocolates (from 0 through 5) was used in both pairs on a given trial. We investigated the effect of having one quantity in common (CQ) in both pairs. Whether the other quantities (OQs) of chocolates were both less than or greater than the CQ, summation still occurred. Accuracy was primarily a function of the ratios of sums to be differentiated. This finding substantiated the earlier conclusion that summation was based on both quantities of each pair and not on some simpler process such as the avoidance of the tray with the smallest single amount or selection of the tray with the single largest amount.     

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.     

Quantity judgments of auditory and visual stimuli by chimpanzees (Pan troglodytes)

Many species can choose between two visual sets of stimuli on the basis of quantity. This is true when sets are both visible, or are presented one set at a time or even one item at a time. However, we know comparatively little about how well nonhuman animals can compare auditory quantities. Here, three chimpanzees (Pan troglodytes) chose between two sets of food items when they only heard each item fall into different containers rather than seeing those items. This method prevented the chimpanzees from summing the amount of visible food they saw because there were no visual cues. Chimpanzees performed well, and their performance matched that of previous experiments with regard to obeying Weber's law. They also performed well with comparisons between a sequentially presented auditory set and a fully visible set, demonstrating that duration of presentation was not being used as a cue. In addition, they accommodated empty sets into these judgments, although not perfectly. Thus, chimpanzees can judge auditory quantities in flexible ways that show many similarities to how they compare visual quantities.     

Colour versus quantity as cues in reverse-reward-competent squirrel monkeys (Saimiri sciureus)

To assess the relative salience of colour and quantity cues, squirrel monkeys previously trained to reach for the smaller of two quantities of food in a reverse-reward contingency task received colour discrimination training. After initial failure to discriminate between two colours of dots under a differential reinforcement regime, they learned the task when the S- colour was associated with zero reward. The monkeys then showed good retention on the original reverse-reward task of 1 versus 4 with pairs of dots presented in S+ or S- colours. However, on "mismatch" trials of 1S- versus 4S+ , only 2 of 4 monkeys tested showed a preference--1 monkey chose based on quantity, the other based on colour. Individual differences and the possible roles of overshadowing and blocking are discussed.     

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.     

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.     

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.     

Differing views: Can chimpanzees do Level 2 perspective-taking?

Although chimpanzees understand what others may see, it is unclear whether they understand how others see things (Level 2 perspective-taking). We investigated whether chimpanzees can predict the behavior of a conspecific which is holding a mistaken perspective that differs from their own. The subject competed with a conspecific over two food sticks. While the subject could see that both were the same size, to the competitor one appeared bigger than the other. In a previously established game, the competitor chose one stick in private first and the subject chose thereafter, without knowing which of the sticks was gone. Chimpanzees and 6-year-old children chose the ‘riskier’ stick (that looked bigger to the competitor) significantly less in the game than in a nonsocial control. Children chose randomly in the control, thus showing Level 2 perspective-taking skills; in contrast, chimpanzees had a preference for the ‘riskier’ stick here, rendering it possible that they attributed their own preference to the competitor to predict her choice. We thus run a follow-up in which chimpanzees did not have a preference in the control. Now, they also chose randomly in the game. We conclude that chimpanzees solved the task by attributing their own preference to the other, while children truly understood the other’s mistaken perspective.     

Ordinal judgments and summation of nonvisible sets of food items by two chimpanzees and a rhesus macaque

Two chimpanzees and a rhesus macaque rapidly learned the ordinal relations between 5 colors of containers (plastic eggs) when all containers of a given color contained a specific number of identical food items. All 3 animals also performed at high levels when comparing sets of containers with sets of visible food items. This indicates that the animals learned the approximate quantity of food items in containers of a given color. However, all animals failed in a summation task, in which a single container was compared with a set of 2 containers of a lesser individual quantity but a greater combined quantity. This difficulty was not overcome by sequential presentation of containers into opaque receptacles, but performance improved if the quantitative difference between sizes was very large.     

Chimpanzees deceive a human competitor by hiding

There is little experimental evidence that any non-human species is capable of purposefully attempting to manipulate the psychological states of others deceptively (e.g., manipulating what another sees). We show here that chimpanzees, one of humans' two closest primate relatives, sometimes attempt to actively conceal things from others. Specifically, when competing with a human in three novel tests, eight chimpanzees, from their first trials, chose to approach a contested food item via a route hidden from the human's view (sometimes using a circuitous path to do so). These findings not only corroborate previous work showing that chimpanzees know what others can and cannot see, but also suggest that when competing for food chimpanzees are skillful at manipulating, to their own advantage, whether others can or cannot see them.     

Summation of symbols by pigeons (Columba livia): the importance of number and mass of reward items

In each of 3 experiments, different sets of 4 pigeons (Columba livia) were trained to discriminate between 2 visual symbols that covered wells containing food items that varied in number, mass, or both. In Experiment 1, the symbols were associated with 0, 1, 3, 5, 7, or 9 pieces of grain reward. The pigeons learned to choose the symbol corresponding to the larger reward, and on summation tests, they chose the pair of symbols that summed to the larger total reward. When number of food pellets was varied but mass of reward was held constant in Experiment 2, preference for the larger number symbols failed to appear. When number was held constant and mass was varied in Experiment 3, the pigeons showed a clear preference for the larger mass symbols on single-symbol and summation tests. These findings show that pigeons summate the value of symbols and are more likely to represent symbols by mass of food reward than by number of food items.     

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.     

Can dogs count?

Numerical competencies have been thoroughly examined in several species, yet relatively few studies have examined such processes in the domestic dog. In an initial experiment, procedures from numerical studies of chimpanzees (0010 and 0015) were adapted for use with 27 domestic dogs. Subjects in these experiments watched as different quantities of food were sequentially dropped into each of two bowls. The subjects were then allowed to select and consume the contents of one of the bowls. Although dogs excelled in a 1 vs 0 condition, their performance failed to significantly surpass chance across all other ratios. In a second experiment with a single subject (a rough collie named Sedona), the procedure was revised so that non-food stimuli were presented simultaneously to the dog on two magnet boards. If Sedona chose the board with the majority of the items, she was rewarded with a piece of food hidden underneath the board. If she made an incorrect choice, she received no reinforcement. Interestingly, Sedona's performance far exceeded that of the dogs in Experiment 1. Implications of these findings for the study of domestic dogs are discussed.     

Chimpanzees do not take into account what others can hear in a competitive situation

Chimpanzees (Pan troglodytes) know what others can and cannot see in a competitive situation. Does this reflect a general understanding the perceptions of others? In a study by Hare et al. (2000) pairs of chimpanzees competed over two pieces of food. Subordinate individuals preferred to approach food that was behind a barrier that the dominant could not see, suggesting that chimpanzees can take the visual perspective of others. We extended this paradigm to the auditory modality to investigate whether chimpanzees are sensitive to whether a competitor can hear food rewards being hidden. Results suggested that the chimpanzees did not take what the competitor had heard into account, despite being able to locate the hiding place themselves by the noise.     

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.     

Quantity judgments of sequentially presented food items by capuchin monkeys (Cebus apella)

Recent assessments have shown that capuchin monkeys, like chimpanzees and other Old World primate species, are sensitive to quantitative differences between sets of visible stimuli. In the present study, we examined capuchins’ performance in a more sophisticated quantity judgment task that required the ability to form representations of food quantities while viewing the quantities only one piece at a time. In three experiments, we presented monkeys with the choice between two sets of discrete homogeneous food items and allowed the monkeys to consume the set of their choice. In Experiments 1 and 2, monkeys compared an entirely visible food set to a second set, presented item-by-item into an opaque container. All monkeys exhibited high accuracy in choosing the larger set, even when the entirely visible set was presented last, preventing the use of one-to-one item correspondence to compare quantities. In Experiment 3, monkeys compared two sets that were each presented item-by-item into opaque containers, but at different rates to control for temporal cues. Some monkeys performed well in this experiment, though others exhibited near-chance performance, suggesting that this species’ ability to form representations of food quantities may be limited compared to previously tested species such as chimpanzees. Overall, these findings support the analog magnitude model of quantity representation as an explanation for capuchin monkeys’ quantification of sequentially presented food items.

Inferences about guessing and knowing by chimpanzees (Pan troglodytes)

The visual perspective-taking ability of 4 chimpanzees (Pan troglodytes) was investigated. The subjects chose between information about the location of hidden food provided by 2 experimenters who randomly alternated between two roles (the guesser and the knower). The knower baited 1 of 4 obscured cups so that the subjects could watch the process but could not see which of the cups contained the reward. The guesser waited outside the room until the food was hidden. Finally, the knower pointed to the correct cup while the guesser pointed to an incorrect one. The chimpanzees quickly learned to respond to the knower. They also showed transfer to a novel variation of the task, in which the guesser remained inside the room and covered his head while the knower stood next to him and watched a third experimenter bait the cups. The results are consistent with the hypothesis that chimpanzees are capable of modeling the visual perspectives of others.     

Rhesus monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) discriminate between knowing and not knowing and collect information as needed before acting

Humans use memory awareness to determine whether relevant knowledge is available before acting, as when we determine whether we know a phone number before dialing. Such metacognition, or thinking about thinking, can improve selection of appropriate behavior. We investigated whether rhesus monkeys (Macaca mulatta) are capable of a simple form of metacognitive access to the contents of short-term memory. Monkeys chose among four opaque tubes, one of which concealed food. The tube containing the reward varied randomly from trial to trial. On half the trials the monkeys observed the experimenter baiting the tube, whereas on the remaining trials their view of the baiting was blocked. On each trial, monkeys were allowed a single chance to select the tube containing the reward. During the choice period the monkeys had the opportunity to look down the length of each tube, to determine if it contained food. When they knew the location of the reward, most monkeys chose without looking. In contrast, when ignorant, monkeys often made the effort required to look, thereby learning the location of the reward before choosing. Looking improved accuracy on trials on which monkeys had not observed the baiting. The difference in looking behavior between trials on which the monkeys knew, and trials on which they were ignorant, suggests that rhesus monkeys discriminate between knowing and not knowing. This result extends similar observations made of children and apes to a species of Old World monkey, suggesting that the underlying cognitive capacities may be widely distributed among primates.     

Do domestic dogs (<Emphasis Type="Italic">Canis lupus familiaris</Emphasis>) perceive the Delboeuf illusion?

In the last decade, visual illusions have been repeatedly used as a tool to compare visual perception among species. Several studies have investigated whether non-human primates perceive visual illusions in a human-like fashion, but little attention has been paid to other mammals, and sensitivity to visual illusions has been never investigated in the dog. Here, we studied whether domestic dogs perceive the Delboeuf illusion. In human and non-human primates, this illusion creates a misperception of item size as a function of its surrounding context. To examine this effect in dogs, we adapted the spontaneous preference paradigm recently used with chimpanzees. Subjects were presented with two plates containing food. In control trials, two different amounts of food were presented in two identical plates. In this circumstance, dogs were expected to select the larger amount. In test trials, equal food portion sizes were presented in two plates differing in size: if dogs perceived the illusion as primates do, they were expected to select the amount of food presented in the smaller plate. Dogs significantly discriminated the two alternatives in control trials, whereas their performance did not differ from chance in test trials with the illusory pattern. The fact that dogs do not seem to be susceptible to the Delboeuf illusion suggests a potential discontinuity in the perceptual biases affecting size judgments between primates and dogs.