Michael J. Beran

Rhesus monkeys (Macaca mulatta) enumerate large and small sequentially presented sets of items using analog numerical representations.

Two rhesus monkeys selected the larger of two sequentially presented sets of items on a computer monitor. In Experiment 1, performance was related to the ratio of set sizes, and the monkeys discriminated between sets with up to 10 items. Performance was not disrupted when 1 set had fewer than 4 items and 1 set had more than 4 items, a critical trial type for differentiating object file and analog models of numerical representation. Experiment 2 controlled the interitem rate of presentation. Experiment 3 included some trials on which number and amount (visual surface area) offered conflicting cues. Experiment 4 varied the total duration of set presentation and the duration of item visibility. In all of the experiments, performance remained high, although total set presentation duration also acted as a partial cue for the monkeys. Overall, the data indicated that rhesus monkeys estimate the approximate number of items in sequentially presented sets and that they are not relying solely on nonnumerical cues such as rate, duration, or cumulative amount.

Chimpanzees use self-distraction to cope with impulsivity

It is unknown whether animals, like humans, can employ behavioural strategies to cope with impulsivity. To examine this question, we tested whether chimpanzees (Pan troglodytes) would use self-distraction as a coping strategy in a situation in which they had to continually inhibit responses to accumulating candies in order to earn a greater amount of those rewards. We tested animals in three conditions in which they were sometimes given a set of toys and were sometimes allowed physical access to the accumulating candies. Chimpanzees allowed the rewards to accumulate longer before responding when they could divert their attention to the toys, and they manipulated the toys more when the candies were physically accessible. Thus, chimpanzees engaged in self-distraction with the toys when such behaviour was most beneficial as a coping mechanism.

Delay of gratification in chimpanzees (Pan troglodytes)

Delay of gratification in 3 chimpanzees (Pan troglodytes) was examined by using a paradigm based on research with children. The chimpanzees either pressed a door-bell button during a trial and received one reward (the immediate reward) or did not press the doorbell and received another reward (the delayed reward). Two chimpanzees were language-trained, and a 3rd was non-language-trained. Foods (one more-preferred and one less-preferred), photographs of those foods, or lexigrams representing those foods were presented to the chimpanzees. All 3 chimpanzees delayed gratification when foods were physically present. One language-trained chimpanzee also delayed gratification with lexigrams present, and the 2nd language-trained chimpanzee delayed gratification in all three conditions. Language competence and early rearing are proposed as explanations for the different performances of these chimpanzees.

Maintenance of self-imposed delay of gratification by four chimpanzees (Pan troglodytes) and an orangutan (Pongo pygmaeus).

Abstract Delay maintenance, which is the continuance over time of the choice to forgo an immediate, less preferred reward for a future, more preferred reward, was examined in 4 chimpanzees (Pan troglodytes) and 1 orangutan (Pongo pygmaeus). In the 1st experiment, the apes were presented with 20 chocolate pieces that were placed, one at a time, into a bowl that was within their reach. The apes could consume the available chocolate pieces at any time during a trial, but no additional pieces would be given. The total length of time taken to place the 20 items into the bowl ranged from 60 s to 180 s. All 5 apes delayed gratification on a majority of trials until all 20 chocolate pieces were presented. Unlike in most experiments with human children using this test situation, attention by the apes to the reward was not detrimental to delay maintenance. In a 2nd experiment with the chimpanzees only, 4 foods of differing incentive value were presented in different trials in the same manner as in Experiment 1. The chimpanzees were highly successful in obtaining all food pieces, and there was no difference in performance as a function of food type.

The Comparative Study of Metacognition: Sharper Paradigms, Safer Inferences

Results that point to animals’ metacognitive capacity bear a heavy burden, given the potential for competing behavioral descriptions. In this article, formal models are used to evaluate the force of these descriptions. One example is that many existing studies have directly rewarded so-called uncertainty responses. Modeling confirms that this practice is an interpretative danger because it supports associative processes and encourages simpler interpretations. Another example is that existing studies raise the concern that animals avoid difficult stimuli not because of uncertainty monitored, but because of aversion given error-causing or reinforcementlean stimuli. Modeling also justifies this concern and shows that this problem is not addressed by the common practice of comparing performance on chosen and forced trials. The models and related discussion have utility for metacognition researchers and theorists broadly, because they specify the experimental operations that will best indicate a metacognitive capacity in humans or animals by eliminating alternative behavioral accounts.

Perception of Food Amounts by Chimpanzees Based on the Number, Size, Contour Length and Visibility of Items.

Nonhuman animals reliably select the largest of two or more sets of discrete items, particularly if those items are food items. However, many studies of these numerousness judgments fail to control for confounds between amount of food e.g., mass or volume) and number of food items. Stimulus dimensions other than number of items also may play a role in how animals perceive sets and make choices. Four chimpanzees (Pan troglodytes) completed a variety of tasks that involved comparisons of food items (graham crackers) that varied in terms of their number, size, and orientation. In Experiment 1, chimpanzees chose between two alternative sets of visible cracker pieces. In Experiment 2, the experimenters presented one set of crackers in a vertical orientation (stacked) and the other in a horizontal orientation. In Experiment 3, the experimenters presented all food items one-at-a-time by dropping them into opaque containers. Chimpanzees succeeded overall in choosing the largest amount of food. They did not rely on number or contour length as cues when making these judgments but instead primarily responded to the total amount of food in the sets. However, some errors reflected choices of the set with the smaller total amount of food but the individually largest single food item. Thus, responses were not optimal because of biases that were not related to the total amount of food in the sets.

Putting the elephant back in the herd: elephant relative quantity judgments match those of other species

The ability to discriminate between quantities has been observed in many species. Typically, when an animal is given a choice between two sets of food, accurate performance (i.e., choosing the larger amount) decreases as the ratio between two quantities increases. A recent study reported that elephants did not exhibit ratio effects, suggesting that elephants may process quantitative information in a qualitatively different way from all other nonhuman species that have been tested (Irie-Sugimoto et al. in Anim Cogn 12:193–199, 2009). However, the results of this study were confounded by several methodological issues. We tested two African elephants (Loxodonta africana) to more thoroughly investigate relative quantity judgment in this species. In contrast to the previous study, we found evidence of ratio effects for visible and nonvisible sequentially presented sets of food. Thus, elephants appear to represent and compare quantities in much the same way as other species, including humans when they are prevented from counting. Performance supports an accumulator model in which quantities are represented as analog magnitudes. Furthermore, we found no effect of absolute magnitude on performance, providing support against an object-file model explanation of quantity judgment.

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.

Delay of gratification and delay maintenance by rhesus macaques (Macaca mulatta).

The authors tested the self-control of rhesus macaques by assessing if they could refrain from reaching into a food container to maximize the accumulation of sequentially delivered food items (a delay-maintenance task). Three different versions of the task varied the quantity and quality of available food items. In the first 2 versions, food items accumulated across the length of the trial until a monkey consumed the items. In the 3rd task, a single less-preferred food item preceded a single more-preferred food item. Some monkeys delayed gratification even with relatively long delays between deliveries of items. However, the data suggested that self-control, in the majority of tested individuals, was not significantly different across different task versions and that self-control by macaques was not as prevalent in these tasks as it is in chimpanzees and human children.

An efficient computerized testing method for the capuchin monkey (Cebus apella): adaptation of the LRC-CTS to a socially housed nonhuman primate species.

Even with advances in automated testing techniques, the capuchin monkey (Cebus apella) can be a difficult species to test in the laboratory, given its social/behavioral tendencies and typical activity pattern. Laboratories that maintain social colonies of capuchin monkeys are able to separate and test individuals, but the process can be very effortful and time consuming, and the resulting data can be modest in quantity. The present article describes procedures and apparatuses that were used to train a colony of computer-naive capuchin monkeys to quickly and reliably isolate themselves from group members and interact with a computerized test system in order to produce a large volume of data. Several elements that were important in motivating the monkeys to participate are discussed.