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To link to full-text access for this article, visit this link: http://dx.doi.org/10.1016/j.anbehav.2007.10.035 Byline: Michael J. Beran, Theodore A. Evans, Emily H. Harris Abstract: 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 judgements 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 influence 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 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 judgements 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. Author Affiliation: Language Research Center, Georgia State University, U.S.A. Article History: Received 16 May 2007; Revised 14 July 2007; Accepted 29 October 2007 Article Note: (miscellaneous) MS. number: A10772R

Metacognition refers to thinking about one’s thinking or knowing what one knows. Research suggests that this ability is not unique to humans and may be shared with nonhuman animals. In particular, great apes have shown behaviors on a variety of tasks that are suggestive of metacognitive ability. Here we combine a metacognitive task, the information-seeking task, with tool use and variable forms of initial information provided to chimpanzees to explore how informational states impact behavioral responses in these apes. Three chimpanzees were presented with an apparatus that contained five locations where food could be hidden. If they pointed to the correct location, they received the reward, but otherwise they did not. We first replicated several existing findings using this method, and then tested novel hypotheses. The chimpanzees were given different types of information across the experiments. Sometimes, they were shown the location of the food reward. Other times, they were shown only one empty location, which was not useful information. The chimpanzees also could use a tool to search any of those locations before making a selection. Chimpanzees typically used the tool to search out the location of the reward when they could not already know where it was, but they did not use the tool when they already had been given that information. One chimpanzee made inferences about the location of hidden food, even when that food was never shown in that location. The final experiment involved hiding foods of differing preference values, and then presenting the chimpanzees with different initial knowledge states (i.e., where the best food was located, where the less-preferred food was located, or where no food was located). All chimpanzees used the tool when they needed to use it to find the best possible item on that trial, but responded by choosing a location immediately when they did not need the tool. This finding highlights that their behavior was not the result of a simple rule following such as pointing to where any food had been seen. [ABSTRACT FROM AUTHOR]

Decision-making largely is influenced by the relative value of choice options, and the value of such options can be determined by a combination of different factors (e.g., the quantity, size, or quality of a stimulus). In this study, we examined the competing influences of quantity (i.e., the number of food items in a set) and quality (i.e., the original state of a food item) of choice items on chimpanzees’ food preferences in a two-option natural choice paradigm. In Experiment 1, chimpanzees chose between sets of food items that were either entirely whole or included items that were broken into pieces before being shown to the chimpanzees. Chimpanzees exhibited a bias for whole food items even when such choice options consisted of a smaller overall quantity of food than the sets containing broken items. In Experiment 2, chimpanzees chose between sets of entirely whole food items and sets of initially whole items that were subsequently broken in view of the chimpanzees just before choice time. Chimpanzees continued to exhibit a bias for sets of whole items. In Experiment 3, chimpanzees chose between sets of new food items that were initially discrete but were subsequently transformed into a larger cohesive unit. Here, chimpanzees were biased to choose the discrete sets that retained their original qualitative state rather than toward the cohesive or clumped sets. These results demonstrate that beyond a food set's quantity (i.e., the value dimension that accounts for maximization in terms of caloric intake), other seemingly non-relevant features (i.e., quality in terms of a set's original state) affect how chimpanzees assign value to their choice options. [ABSTRACT FROM AUTHOR]

Prospective memory is remembering to do something at a future time. A growing body of research supports that prospective memory may exist in nonhuman animals, but the methods used to test nonhuman prospective memory differ from those used with humans. The current work tests prospective memory in chimpanzees using a method that closely approximates a typical human paradigm. In these experiments, the prospective memory cue was embedded within an ongoing task. Tokens representing food items could be used in one of two ways: in a matching task with pictures of items (the ongoing task) or to request a food item hidden in a different location at the beginning of the trial. Chimpanzees had to disengage from the ongoing task in order to use the appropriate token to obtain a higher preference food item. In , chimpanzees effectively matched tokens to pictures, when appropriate, and disengaged from the ongoing task when the token matched the hidden item. In , performance did not differ when the target item was either hidden or visible. This suggested no effect of cognitive load on either the prospective memory task or the ongoing task, but performance was near ceiling, which may have contributed to this outcome. In , we created a more challenging version of the task. More errors on the matching task occurred before the prospective memory had been carried out, and this difference seemed to be limited to the hidden condition. This finding parallels results from human studies and suggests that working memory load and prospective memory may have a similar relationship in nonhuman primates. [ABSTRACT FROM AUTHOR]

Token exchange inherently introduces an element of delay between behavior and reward and so token studies may help us better understand delay of gratification and self-control. To examine this possibility, we presented three language-trained chimpanzees with repeated choices involving different foods that could be eaten immediately or lexigram (graphic symbol) tokens that represented (and could be traded for) foods later. When both options were foods, chimpanzees always chose more preferred foods over less preferred foods. When both options were lexigram tokens representing those same foods, performance remained the same as chimpanzees selected the higher value token and then traded it for food. Then, when faced with choosing a token that could be traded later or choosing a food item that could be eaten immediately, most chimpanzees learned to make whatever response led to the more preferred food. They did this even when that meant selecting a high value lexigram token that could be traded only 2 to 3 min later instead of a medium value, but immediately available, food item. Thus, chimpanzees flexibly selected tokens even though such selections necessarily delayed gratification and required forgoing immediately available food. This finding illustrates the utility of symbolic token exchange for assessing self-control in nonhuman animals. Am. J. Primatol. 74:864-870, 2012. © 2012 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Self-control is often required in natural situations involving interactions with other individuals, and personal self-control can be compromised if other individuals act impulsively. In this study, we tested self-control in pairs of chimpanzees in a variety of settings where at least one chimpanzee of each pair performed an established test for self-control in which candies accumulated one at time as long as the chimpanzee did not eat any of them. When tested alone, some chimpanzees exhibited greater self-control as compared to when tested alongside a chimpanzee that independently performed the same type of test. However, when the nonfocal animal freely consumed rewards while the focal chimpanzee performed the accumulation task, the self-control of some focal chimpanzees was elevated as compared to when working alone. Finally, when the focal and nonfocal animals worked jointly on the same test and the number of rewards accumulated was dependent on both animals' continued ability to inhibit eating the items, chimpanzees performed the same when housed together or in adjacent enclosures. On the whole, the effects of social setting were modest, but these results may relate to the literature on vicarious depletion of self-control, and they present interesting avenues for future research. [ABSTRACT FROM AUTHOR]

We presented four chimpanzees with a series of tasks that involved comparing two token sets or comparing a token set to a quantity of food. Selected tokens could be exchanged for food items on a one-to-one basis. Chimpanzees successfully selected the larger numerical set for comparisons of 1 to 5 items when both sets were visible and when sets were presented through one-by-one addition of tokens into two opaque containers. Two of four chimpanzees used the number of tokens and food items to guide responding in all conditions, rather than relying on token color, size, total amount, or duration of set presentation. These results demonstrate that judgments of simultaneous and sequential sets of stimuli are made by some chimpanzees on the basis of the numerousness of sets rather than other non-numerical dimensions. The tokens were treated as equivalent to food items on the basis of their numerousness, and the chimpanzees maximized reward by choosing the larger number of items in all situations. [ABSTRACT FROM AUTHOR]

The authors investigated choice behavior by chimpanzees in five experiments involving choices between different amounts of food. Chimpanzees did not maximize the amount of food they obtained when choosing between a single 20-g banana piece and another option containing a 20-g piece and a 5-g piece. This was true even though they successfully discriminated between 20-g and 25-g banana pieces in other trials. When items in the mixed option were stacked, however, the chimpanzees chose the larger amount. Later experiments indicated that changing the magnitude of the two amounts did not change performance if the difference in magnitude between the two options remained the same (e.g., 40 g plus 10 g vs. 40 g). However, chimpanzees did improve when the two-item option was increased in its magnitude relative to the single slice (e.g., 20 g plus 10 g vs. 20 g). These results indicated that chimpanzees undervalued the total amount of food in sets when items differed in size and did not appear to be whole. Another experiment confirmed that it was this notion of wholeness that evoked suboptimal responding because chimpanzees were successful in the same comparisons with a different type of food that appeared less fractionated when presented as two pieces. These results provide evidence of suboptimal responding in some natural choice situations that prevents chimpanzees from maximizing food intake. [ABSTRACT FROM AUTHOR]

Abstract: We tested four chimpanzees in a self-control task in which food rewards accumulated as long as they were not eaten. In one condition, the chimpanzees had to perform a computer task that directly led to the delivery of the food rewards. In another condition, working on the computerized task was not required and any such work was not linked to the delivery of rewards. The third condition offered no computerized task (chimpanzees simply waited for food rewards to be delivered). Three of four chimpanzees showed no effect of the work scenario on delay of gratification. The one chimpanzee that showed an influence of work scenario on self-control was the overall poorest performing animal. This animal delayed gratification the longest, however, when work was required and reward delivery was directly linked to that work. Therefore, although there is little evidence linking delay of gratification to work requirements in chimpanzees, chimpanzees with lower overall self-control might benefit from having some work available if reward accumulation is contingent on performing that work. [Copyright &y& Elsevier]

Abstract: Previous research in our laboratory has demonstrated that chimpanzees can delay gratification by inhibiting consumption of available food items for as long as 3min as an experimenter transfers additional food items from a transparent container to a bowl placed in front of the subject. In this study, we examined the influence of the visibility of the food source, as well as the presence of the experimenter, on four chimpanzees’ self-control in this paradigm. In Experiment 1 an experimenter transferred 15 preferred food items between a distant opaque container and a bowl placed in front of the subject. In Experiment 2 we tested the chimpanzees with an automated system that (in the absence of the experimenter) transferred up to 36 highly preferred food items from a universal food dispenser to a container located either inside or outside of the subject''s enclosure. There were no differences in self-directed behaviors or attentiveness to the food items between the self-imposed and externally imposed delay conditions. A final experiment with the automated paradigm indicated that individuals could delay gratification for up to 11min in order to obtain all 36 food items. [Copyright &y& Elsevier]

Abstract: In three experiments, four chimpanzees made choices between two visible food options to assess the validity of the selective value effect (the assignment of value to only the most preferred type of food presented in a comparison). In Experiment 1, we established that all chimpanzees preferred single banana pieces to single apple pieces before presenting the critical test. In this test two chimpanzees preferred a mix of one banana piece and one apple piece to a single banana piece when both banana pieces were approximately the same size, but two chimpanzees were indifferent between the two options, exhibiting the selective value effect. In Experiment 2, when the banana pieces in both options were more closely equated in size, the chimpanzees were biased to choose the single banana piece over the mixed array even though this was the smaller total amount of food. However, in Experiment 3, when we introduced longer intervals between each trial, the chimpanzees preferred the mixed set and thus the larger total amount of food. The results demonstrate that only some chimpanzees exhibit the choice pattern indicative of the selective value effect, and they do so only when item size is not carefully controlled and trials are presented quickly in succession. Thus, the behavior pattern originally labeled the selective value effect may actually be explained by a combination of chimpanzees’ sensitivity to small differences in preferred food amount and chimpanzees tendency to avoid less preferred foods that would delay the acquisition of further preferred food items. [Copyright &y& Elsevier]

Three chimpanzees performed a computerized memory task in which auditory feedback about the accuracy of each response was delayed. The delivery of food rewards for correct responses also was delayed and occurred in a separate location from the response. Crucially, if the chimpanzees did not move to the reward-delivery site before food was dispensed, the reward was lost and could not be recovered. Chimpanzees were significantly more likely to move to the dispenser on trials they had completed correctly than on those they had completed incorrectly, and these movements occurred before any external feedback about the outcome of their responses. Thus, chimpanzees moved (or not) on the basis of their confidence in their responses, and these confidence movements aligned closely with objective task performance. These untrained, spontaneous confidence judgments demonstrated that chimpanzees monitored their own states of knowing and not knowing and adjusted their behavior accordingly. [ABSTRACT FROM AUTHOR]

Human and nonhuman primates are not mentally constrained to the present. They can remember the past and--at least to an extent--anticipate the future. Anticipation of the future ranges from long-term prospection such as planning for retirement to more short-term future-oriented cognition such as planning a route through a maze. Here we tested a great ape species (chimpanzees), an Old World monkey species (rhesus macaques), a New World monkey species (capuchin monkeys), and human children on a computerized maze task. All subjects had to move a cursor through a maze to reach a goal at the bottom of the screen. For best performance on the task, subjects had to "plan ahead" to the end of the maze to move the cursor in the correct direction, avoid traps, and reverse directions if necessary. Mazes varied in difficulty. Chimpanzees were better than both monkey species, and monkeys showed a particular deficit when moving away from the goal or changing directions was required. Children showed a similar pattern to monkeys regarding the effects of reversals and moves away from the goal, but their overall performance in terms of correct maze completion was similar to the chimpanzees. The results highlight similarities as well as differences in planning across species and the role that inhibitory control may play in future-oriented cognition in primates. [ABSTRACT FROM AUTHOR]

Nonhuman animals quantify all manner of things, and the way in which this is done is fairly well understood. However, little research has been conducted to determine how they know what is or is not relevant in the instances in which they quantify stimuli. We assessed how four chimpanzees chose between two sets of food items when the items were distributed across separate spatial arrays. Each item was covered by a container, and then was revealed in sequence so that neither whole set was visible at one time. After all containers were revealed, some were revealed again. The chimpanzees should have ignored items that were seen a second time and instead enumerated each item only once. In another test, some of the items were transposed in location and then uncovered again. Here, the chimpanzees needed to recognize that the newly shown food items were ones they already had seen. Overall, the chimpanzees were successful in selecting the truly larger array of items despite these potential distracting re-presentations of items. Discrimination performance also reflected analogue magnitude estimation because comparisons of sets that differed by larger amounts were easier than comparisons that differed by smaller amounts. Thus, chimpanzee quantity judgments for nonvisible sets of items are inexact, but they include an aspect of control for determining when items are uniquely presented versus re-presented. [ABSTRACT FROM AUTHOR]

Abstract: Prospective memory involves the encoding, retention, and implementation of an intended future action. Although humans show many forms of prospective memory, less is known about the future oriented processes of nonhuman animals, or their ability to use prospective memory. In this experiment, a chimpanzee named Panzee, who had learned to associate geometric forms called lexigrams with real-world referents, was given a prospective memory test. Panzee selected between two foods the one she wanted to receive more immediately. That food was scattered in an outdoor yard where she could forage for it. Also outdoors were lexigram tokens, one of which represented the food item that remained indoors throughout a 30min period, and that could be obtained if Panzee brought in the token that matched that food item. After foraging for the selected food item, Panzee consistently remembered to retrieve and return the correct token when food was available indoors, whereas on control trials involving no indoor food she rarely returned a token. This indicated that Panzee encoded information relevant to the future action of token retrieval after extended delays for one type of food, even when a more immediately preferred food was available. [Copyright &y& Elsevier]