Maintenance of Self-Imposed Delay of Gratification by Four Chimpanzees (Pan troglodytes) and an Orangutan (Pongo pygmaeus)

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.     

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

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.     

Reverse-reward learning in squirrel monkeys (Saimiri sciureus): retesting after 5 years, and assessment on qualitative transfer

Seven squirrel monkeys (Saimiri sciureus) previously trained on reverse-reward tasks were presented with the original "1-versus-4" task after a 5-year interval without reverse-reward experience (Experiment 1). None of them reliably selected the smaller food array; however, at around chance level, their performance was superior to when they were first exposed to the task almost 6 years previously, suggesting some long-term memory retention. One naive monkey consistently selected the larger array, as expected. In Experiment 2, trials consisting of 1 versus 1 piece of two qualitatively different types of food were interspersed among familiar 1-versus-4 trials. None of five monkeys tested reliably selected the less-preferred food to get the more preferred food as the reward, and one monkey scored below chance. However, when one piece of low-preference food was paired with four pieces of high-preference food (Experiment 3), all four monkeys tested avoided reaching for the latter and thereby obtained it as the reward; two monkeys obtained perfect scores on these trials. These two monkeys were trained on a specific qualitative reverse-reward pairing and then again tested on new pairings (Experiment 4), but transfer was incomplete. Compound trials that pit quantity against quality in novel ways appear taxing for squirrel monkeys, despite competence in reverse-reward on both dimensions separately.     

Token reinforcement, choice, and self-control in pigeons.

Pigeons were exposed to self-control procedures that involved illumination of light-emitting diodes (LEDs) as a form of token reinforcement. In a discrete-trials arrangement, subjects chose between one and three LEDs; each LED was exchangeable for 2-s access to food during distinct posttrial exchange periods. In Experiment 1, subjects generally preferred the immediate presentation of a single LED over the delayed presentation of three LEDs, but differences in the delay to the exchange period between the two options prevented a clear assessment of the relative influence of LED delay and exchange-period delay as determinants of choice. In Experiment 2, in which delays to the exchange period from either alternative were equal in most conditions, all subjects preferred the delayed three LEDs more often than in Experiment-1. In Experiment 3, subjects preferred the option that resulted in a greater amount of food more often if the choices also produced LEDs than if they did not. In Experiment 4, preference for the delayed three LEDs was obtained when delays to the exchange period were equal, but reversed in favor of an immediate single LED when the latter choice also resulted in quicker access to exchange periods. The overall pattern of results suggests that (a) delay to the exchange period is a more critical determinant of choice than is delay to token presentation; (b) tokens may function as conditioned reinforcers, although their discriminative properties may be responsible for the self-control that occurs under token reinforcer arrangements; and (c) previously reported differences in the self-control choices of humans and pigeons may have resulted at least in part from the procedural conventions of using token reinforcers with human subjects and food reinforcers with pigeon subjects.     

A reversed-reward contingency task reveals causal knowledge in chimpanzees (Pan troglodytes)

In the reversed-reward contingency task, subjects are required to choose the less preferred of two options in order to obtain the more preferred one. Usually, this task is used to measure inhibitory skills, but it could also be used to measure how strong the subjects’ preferences are. We presented chimpanzees with support tasks where only one of two paper strips could physically bring food into reach. Subjects were rewarded for choosing the non-functional strip. In Experiment 1, subjects failed to pick the non-baited strip. In Experiment 2, subjects failed to pick the broken strip. Chimpanzees performed worse in these tasks than in other similar tasks where instead of paper strips, there were similar shapes painted on a platform. The fact that subjects found the reversed-reward contingency task based on causality more difficult to solve than a perceptually similar task with no causality involved (i.e., arbitrary) suggests that they did not treat real strips as an arbitrary task. Instead, they must have had some causal knowledge of the support problem that made them prefer functional over non-functional strips despite the contrary reward regime.     

Chimpanzees (Pan troglodytes) conceal visual and auditory information from others

Chimpanzees (Pan troglodytes) competed with a human for food. The human sat inside a booth, with 1 piece of food to her left and 1 to her right, which she could retract from her chimpanzee competitor's reach as needed. In Experiment 1, chimpanzees could approach either side of the booth unseen but then had to reach through 1 of 2 tunnels (1 clear, 1 opaque) for the food. In Experiment 2, both tunnels were clear and the human was looking away, but 1 of the tunnels made a loud noise when it was opened. Chimpanzees preferentially reached through the opaque tunnel in the first study and the silent tunnel in the second, successfully concealing their taking of the food from the human competitor in both cases. These results suggest that chimpanzees can, in some circumstances, actively manipulate the visual and auditory perception of others by concealing information from them.     

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.     

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).     

Referential communication by chimpanzees (Pan troglodytes)

Two experiments were conducted to assess the referential function of chimpanzee (Pan troglodytes) gestures to obtain food. The chimpanzees received 1 trial per condition. In Experiment 1 (N = 101), in full view of the chimpanzee, a banana was placed on top of 1 of 2 inverted buckets or was hidden underneath 1 of the buckets. In Experiment 2 (N = 35), 4 conditions were presented in constant order: (a) no food, no observer; (b) no food, observer present; (c) food present, no observer; and (d) food present, observer present. Gestures and visual orienting were used socially and referentially. The capacity for nonverbal reference may predate the Hominidae-Pongidae split, and the development of nonverbal reference may be independent of human species-specific adaptations for speech.     

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.     

Token transfer between mother and offspring chimpanzees (Pan troglodytes): mother–offspring interaction in a competitive situation

Chimpanzees can flexibly use tokens in cognitive tasks, but it is still unknown if they can share and/or compete over tokens as they do for food. This study aimed to evaluate the interactions spontaneously occurring between mother and offspring chimpanzees when tokens exchangeable for food were provided. Forty tokens were scattered on the floor in an experimental playroom. Three mother and offspring chimpanzee pairs were tested. Each token was exchangeable for a piece of food in a vending machine installed on the wall of the playroom. In the beginning of the study, both mother and offspring took tokens and exchanged them for pieces of food independently. Later, two offspring started to take more tokens than their mothers. At that time, the offspring whimpered or cried more often than during earlier sessions. This behavior compelled the mothers to abstain from taking tokens. The mothers sometimes shared their tokens with their offspring, or were tolerant of their offspring taking their tokens from their hand. For one pair, the offspring sometimes shared tokens with her mother when her mother begged for the tokens. These results suggest that chimpanzees’ cognitive abilities enable them not only to use tokens, but also to compete for tokens, as they do for food. The results also suggest that token sharing between mother and offspring may be bidirectional and that transfer of tokens mainly occurs as a result of begging, although on some occasion offspring were able to obtain a token directly from his/her mother through tolerated scrounging.     

Scaling reward value with demand curves versus preference tests

In Experiment 1, six capuchins lifted a weight during a 10-min session to receive a food piece. Across conditions, the weight was increased across six different amounts for three different food types. The number of food pieces obtained as a function of the weight lifted was fitted by a demand equation that is hypothesized to quantify food value. For most subjects, this analysis showed that the three food types differed little in value. In Experiment 2, these monkeys were given pairwise choices among these food types. In 13 of 18 comparisons, preferences at least equaled a 3-to-1 ratio; in seven comparisons, preference was absolute. There was no relation between values based on degree of preference versus values based on the demand equation. When choices in the present report were compared to similar data with these subjects from another study, between-study lability in preference emerged. This outcome contrasts with the finding in demand analysis that test–retest reliability is high. We attribute the unreliability and extreme assignment of value based on preference tests to high substitutability between foods. We suggest use of demand analysis instead of preference tests for studies that compare the values of different foods. A better strategy might be to avoid manipulating value by using different foods. Where possible, value should be manipulated by varying amounts of a single food type because, over an appropriate range, more food is consistently more valuable than less. Such an approach would be immune to problems in between-food substitutability.     

Spatial memory and monitoring of hidden items through spatial displacements by chimpanzees (Pan troglodytes)

This study examined chimpanzee (Pan troglodytes) short-term memory for food location in near space. In Experiments 1 and 2, either 1 or 2 items (chocolate pieces) were hidden in an array of 3 or 5 containers that either remained stationary or were rotated 180 degrees or 360 degrees. When the array remained stationary, the chimpanzees remembered both item locations. When arrays were rotated, however, chimpanzees found only 1 item. In Experiment 3, 2 items were hidden in an array of 7 cups. Both items were found at levels significantly better than chance. Ninety percent of errors were made after the 1st item was found, and errors reflected memory failure rather than a failure of inhibitory control.     

Self-control and impulsiveness in children and adults: Effects of food preferences

Experiment 1 used 6 preschool boys and Experiment 2 used 6 adult women to explore the effects of food preference on humans' choice in self-control paradigms. The boys showed a higher proportion of responses for more delayed, larger reinforcers (a measure of self-control) when those choices resulted in receipt of the most preferred food compared to when those choices resulted in the least preferred food. Further, the boys chose the less delayed, smaller reinforcers significantly more often when only those choices, as opposed to both choices, resulted in the most preferred food. Conversely, they chose the more delayed, larger reinforcers significantly more often when only those choices, as opposed to both choices, resulted in the most preferred food. Finally, the women demonstrated significantly less sensitivity to reinforcer amount relative to sensitivity to reinforcer delay (another measure of self-control) when they had a higher preference for the juice received as the less delayed, smaller reinforcer than for the juice received as the more delayed, larger reinforcer. Together, the results show that subjects' food preferences can influence self-control for food reinforcers.     

Time preferences in long-tailed macaques (Macaca fascicularis) and humans (Homo sapiens)

Rosati et al. (Curr Biol 17(19):1663–1668, 2007) found in a self-control test in which choice was between a smaller, immediately delivered food and a larger, delayed food, that chimpanzees preferred the larger reward (self-control); humans, however, preferred the smaller reward (impulsivity). They attributed their results to a species difference in self-control. In Experiment 1, monkeys (long-tailed macaques) were exposed to a self-control task in two conditions: where the food was hidden under differently colored bowls and where it was visible. When these two conditions were compared, choice shifted from greater preference for the impulsive alternative in the hidden condition to greater preference for the self-control alternative in the visible condition. Additionally, in both conditions, preference shifted from self-control to impulsivity over sessions. These results were explained in terms of the reversed-contingency effect (a propensity to reach for more over less when rewards are visible) and not to a capacity for self-control. In Experiment 2, humans that demonstrated preference for more over less in choice preferred the impulsive alternative when choice to either alternative was followed by the same intertrial interval—a preference that accelerates trial rates relative to preference of the self-control alternative. When trial rates were equated so that neither choice accelerated session’s end, humans demonstrated self-control. These results suggest that Rosati et al.’s demonstration of impulsivity in humans was due to participants’ desire to minimize session time.     

Grip morphology and hand use in chimpanzees (Pan troglodytes): evidence of a left hemisphere specialization in motor skill

Three experiments on grip morphology and hand use were conducted in a sample of chimpanzees. In Experiment 1, grip morphology when grasping food items was recorded, and it was found that subjects who adopted a precision grip were more right-handed than chimpanzees using other grips. In Experiment 2, the effect of food type on grasping was assessed. Smaller food items elicited significantly more precision grips for the right hand. In Experiment 3, error rates in grasping foods were compared between the left and right hands. Significantly more errors were made for the left compared with the right hand. The cumulative results indicate that chimpanzees show a left-hemisphere asymmetry in motor skill that is associated with the use of precision grips.     

Stock optimizing in choice when a token deposit is the operant.

Each of 2 monkeys typically earned their daily food ration by depositing tokens in one of two slots. Tokens deposited in one slot dropped into a bin where they were kept (token kept). Deposits to a second slot dropped into a bin where they could be obtained again (token returned). In Experiment 1, a fixed-ratio (FR) 5 schedule that provided two food pellets was associated with each slot. Both monkeys preferred the token-returned slot. In Experiment 2, both subjects chose between unequal FR schedules with the token-returned slot always associated with the leaner schedule. When the FRs were 2 versus 3 and 2 versus 6, preferences were maintained for the token-returned slot; however, when the ratios were 2 versus 12, preference shifted to the token-kept slot. In Experiment 3, both monkeys chose between equal-valued concurrent variable-interval variable-interval schedules. Both monkeys preferred the slot that returned tokens. In Experiment 4, both monkeys chose between FRs that typically differed in size by a factor of 10. Both monkeys preferred the FR schedule that provided more food per trial. These data show that monkeys will choose so as to increase the number of reinforcers earned (stock optimizing) even when this preference reduces the rate of reinforcement (all reinforcers divided by session time).     

Keeping track of time: evidence for episodic-like memory in great apes

Episodic memory, as defined by Tulving, can be described in terms of behavioural elements (what, where and when information) but it is also accompained by an awareness of one’s past (chronesthesia) and a subjective conscious experience (autonoetic awareness). Recent experiments have shown that corvids and rodents recall the where, what and when of an event. This capability has been called episodic-like memory because it only fulfils the behavioural criteria for episodic memory. We tested seven chimpanzees, three orangutans and two bonobos of various ages by adapting two paradigms, originally developed by Clayton and colleagues to test scrub jays. In Experiment 1, subjects were fed preferred but perishable food (frozen juice) and less preferred but non-perishable food (grape). After the food items were hidden, subjects could choose one of them either after 5 min or 1 h. The frozen juice was still available after 5 min but melted after 1 h and became unobtainable. Apes chose the frozen juice significantly more after 5 min and the grape after 1 h. In Experiment 2, subjects faced two baiting events happening at different times, yet they formed an integrated memory for the location and time of the baiting event for particular food items. We also included a memory task that required no temporal encoding. Our results showed that apes remember in an integrated fashion what, where and when (i.e., how long ago) an event happened; that is, apes distinguished between different events in which the same food items were hidden in different places at different times. The temporal control of their choices was not dependent on the familiarity of the platforms where the food was hidden. Chimpanzees’ and bonobos’ performance in the temporal encoding task was age-dependent, following an inverted U-shaped distribution. The age had no effect on the performance of the subjects in the task that required no temporal encoding.     

Numerical judgments by chimpanzees (Pan troglodytes) in a token economy

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.     

Do chimpanzees know what others can and cannot do? Reasoning about ‘capability’

Much recent comparative work has been devoted to exploring what nonhuman primates understand about physical causality. However, few laboratory experiments have attempted to test what nonhumans understand about what physical acts others are capable of performing. We tested seven chimpanzees’ ability to predict which of two human experimenters could deliver a tray containing a food reward. In the ‘floor’ condition, legs were required to push the tray toward the subject. In the ‘lap’ condition, arms were required to hand the tray to the subject. In Exp. 1, chimpanzees begged (by gesturing) to either an experimenter whose legs were not visible (LNV) or whose arms were not visible (ANV). Rather than flexibly altering their preferences between conditions, the chimpanzees preferred the ANV experimenter regardless of the task. In subsequent experiments, we manipulated various factors that might have controlled the chimpanzees’ preferences, such as (a) distance between experimenter and subject (Experiment 2), (b) amount of occlusion of experimenters’ body (Experiments 2 and 3), (c) contact with the food tray (Experiments 3 and 4) and (d) positioning of barriers that either impeded the movement of the limbs or not (Experiment 5). The chimpanzees’ performance was best explained by attention to cues such as perceived proximity, contact, and maximal occlusion of body that although highly predictive in certain tasks, were irrelevant in others. When the discriminative role of such cues was eliminated, performance fell to chance levels, indicating that chimpanzees do not spontaneously (or after considerable training) use limb visibility as a cue to predict the ability of a human to perform particular physical tasks. Thus, the current findings suggest a possible failure of causal reasoning in the context of reasoning about the use of the limbs to perform physical acts.