Theodore A. Evans

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.

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.

Self-control and tool use in tufted capuchin monkeys (Cebus Apella)

Self-control is defined as forgoing immediate gratification to obtain a greater reward. Tool use may relate to self-control because both behaviors may require foresight and deliberate control over ones actions. The authors assessed 20 capuchin monkeys (Cebus apella) for the ability to delay gratification in a tool task. Subjects were given rod-shaped food items that could either be consumed immediately or be carried to an apparatus and used to extract a more preferred food. The authors found that some monkeys were able to exhibit self-control. Monkeys with relatively more tool use experience demonstrated the greatest levels of self-control. These results indicate that capuchins are capable of delaying gratification when a higher quality reinforcer is present and that tool experience can influence levels of self-control in this task.

Delaying gratification for food and tokens in capuchin monkeys (Cebus apella) and chimpanzees (Pan troglodytes): when quantity is salient, symbolic stimuli do not improve performance

Capuchin monkeys have been tested for the capacity to delay gratification for accumulating rewards in recent studies and have exhibited variable results. Meanwhile, chimpanzees have consistently excelled at this task. However, neither species have ever been tested at accumulating symbolic tokens instead of food items, even though previous reports indicate that tokens sometimes facilitate performance in other self-control tasks. Thus, in the present study, we tested capuchin monkeys and chimpanzees for their capacity to delay gratification in a delay maintenance task, in which an experimenter presented items, one at a time, to within reach of an animal for as long as the animal refrained from taking them. In Experiment 1, we assessed how long capuchin monkeys could accumulate items in the delay maintenance task when items were food rewards or tokens exchangeable for food rewards. Monkeys accumulated more food rewards than they did tokens. In Experiment 2, we tested capuchin monkeys and chimpanzees in a similar accumulation test. Whereas capuchins again accumulated more food than tokens, all chimpanzees but one showed no difference in performance in the two conditions. These findings provide additional evidence that chimpanzees exhibit greater self-control capacity in this task than do capuchin monkeys and indicate that symbolic stimuli fail to facilitate delay maintenance when they do not abstract away from the quantitative dimension of the task. This is consistent with previous findings on the effects of symbols on self-control and illuminates what makes accumulation a particularly challenging task.

Auditory--visual cross-modal perception of communicative stimuli in tufted capuchin monkeys (Cebus apella).

Research on cross-modal performance in nonhuman primates is limited to a small number of sensory modalities and testing methods. To broaden the scope of this research, the authors tested capuchin monkeys (Cebus apella) for a seldom-studied cross-modal capacity in nonhuman primates, auditory-visual recognition. Monkeys were simultaneously played 2 video recordings of a face producing different vocalizations and a sound recording of 1 of the vocalizations. Stimulus sets varied from naturally occurring conspecific vocalizations to experimentally controlled human speech stimuli. The authors found that monkeys preferred to view face recordings that matched presented vocal stimuli. Their preference did not differ significantly across stimulus species or other stimulus features. However, the reliability of the latter set of results may have been limited by sample size. From these results, the authors concluded that capuchin monkeys exhibit auditory-visual cross-modal perception of conspecific vocalizations.

Delay Choice Versus Delay Maintenance: Different Measures of Delayed Gratification in Capuchin Monkeys (Cebus apella)

Delaying gratification involves 2 components: (1) delay choice (selecting a delayed reward over an immediate one) and (2) delay maintenance (sustaining the decision to delay gratification even if the immediate reward is available during the delay). Two tasks most commonly have explored these components in primates: the intertemporal choice task and the accumulation task. It is unclear whether these tasks provide equivalent measures of delay of gratification. Here, we compared the performance on the intertemporal choice task and the accumulation task of capuchin monkeys (Cebus apella) belonging to 2 study populations. We found only limited evidence of a significant correlation in performance. Consequently, in contrast to what is often assumed, our data provide only partial support for the hypothesis that these tasks provide equivalent measures of delay of gratification.

Discrimination reversal learning in capuchin monkeys (Cebus apella).

Learning styles in capuchin monkeys were assessed with a computerized reversal-learning task called the mediational paradigm. First, monkeys were trained to respond with 90% accuracy on a two-choice discrimination (A+B-−). Then the authors examined differences in performance on three different types of reversal trials (A-−B+, A-−C+, B+C-−), each of which offered differing predictions for performance, depending on whether the monkeys were using associative cues or rule-based strategies. Performance indicated that the monkeys mainly learned to avoid the B stimulus during training, as the A-−C+ condition produced the best performance levels. Therefore, negative stimuli showed greater control over responding after reversal and reflected a more associative rather than rule-based form of learning.

Capuchin monkeys (Cebus apella) let lesser rewards pass them by to get better rewards

Self-control is defined as foregoing an immediate reward to gain a larger delayed reward. Methods used to test self-control comparatively include inter-temporal choice tasks, delay of gratification tasks, and accumulation tasks. To date, capuchin monkeys have shown different levels of self-control across tasks. This study introduced a new task that could be used comparatively to measure self-control in an intuitive context that involved responses that required no explicit training. Capuchin monkeys (Cebus apella) were given a choice between two food items that were presented on a mechanized, revolving tray that moved those foods sequentially toward the monkeys. A monkey could grab the first item or wait for the second, but was only allowed one item. Most monkeys in the study waited for a more highly preferred food item or a larger amount of the same food item when those came later, and they inhibited the prepotent response to grab food by not reaching out to take less-preferred foods or smaller amounts of food that passed directly in front of them first. These data confirm that the mechanisms necessary for self-control are present in capuchin monkeys and indicate that the methodology can be useful for broader comparative assessments of self-control.