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.     

Behavioural development in a matching-to-sample task and token use by an infant chimpanzee reared by his mother

We investigated the behavioural and cognitive development of a captive male infant chimpanzee, Ayumu, raised by his mother, Ai. Here we report Ayumu's achievements up to the age of 2 years and 3 months, in the context of complex computer-controlled tasks. From soon after birth, Ayumu had been present during an experiment performed by his mother. The task consisted of two phases, a matching-to-sample task in which she received token rewards, and the insertion of these tokens into a vending machine to obtain food rewards. Ayumu himself received no reward or encouragement from humans for any of the actions he exhibited during the experiment. At the age of 9 months and 3 weeks, Ayumu performed his first matching-to-sample trial. At around 1 year and 3 months, he began to perform them consistently. Also during this period, he frequently stole food rewards from his mother. At 2 years and 3 months, Ayumu succeeded for the first time in inserting a token into the vending machine. Once he had succeeded in using a token, he performed both phases of the task in sequence 20 times consecutively. The infant's behaviour was not shaped by food rewards but by a strong motivation to copy his mother's behaviour. Our observations of Ayumu thus mirror the learning processes shown by wild chimpanzees.     

Social comparison mediates chimpanzees’ responses to loss,not frustration

Why do chimpanzees react when their partner gets a better deal than them? Do they note the inequity or do their responses reflect frustration in response to unattainable rewards? To tease apart inequity and contrast, we tested chimpanzees in a series of conditions that created loss through individual contrast, through inequity, or by both. Chimpanzees were tested in four social and two individual conditions in which they received food rewards in return for exchanging tokens with an experimenter. In conditions designed to create individual contrast, after completing an exchange, the chimpanzees were given a relatively less-preferred reward than the one they were previously shown. The chimpanzees’ willingness to accept the less-preferred rewards was independent of previously offered foods in both the social and individual conditions. In conditions that created frustration through inequity, subjects were given a less-preferred reward than the one received by their partner, but not in relation to the reward they were previously offered. In a social context, females were more likely to refuse to participate when they received a less-preferred reward than their partner (disadvantageous inequity), than when they received a more-preferred reward (advantageous inequity). Specifically, the females’ refusals were typified by refusals to exchange tokens rather than refusals to accept food rewards. Males showed no difference in their responses to inequity or individual contrast. These results support previous evidence that some chimpanzees’ responses to inequity are mediated more strongly by what others receive than by frustration effects.     

Bonobos and orangutans,but not chimpanzees,flexibly plan for the future in a token-exchange task

Non-human animals, including great apes, have been suggested to share some of the skills for planning that humans commonly exhibit. A crucial difference between human and non-human planning may relate to the diversity of domains and needs in which this skill is expressed. Although great apes can save tools for future use, there is little evidence yet that they can also do so in other contexts. To investigate this question further, we presented the apes with a planning token-exchange task that differed from standard tool-use tasks. Additionally, we manipulated the future outcome of the task to investigate planning flexibility. In the Exchange condition, subjects had to collect, save and transport tokens because they would need them 30 min later to exchange them for food with a human, i.e., “bring-back” response. In the Release condition, the collection and transport of tokens were not needed as no exchange took place after 30 min. Out of 13 subjects, eight solved the task at least once in the Exchange condition, with chimpanzees appearing less successful than the other species. Importantly, three individuals showed a clear differential response between conditions by producing more “bring-back” responses in the Exchange than in the Release conditions. Those bonobo and orangutan individuals hence adapted their planning behavior according to changing needs (i.e., they brought tokens back significantly more often when they would need them). Bonobos and orangutans, unlike chimpanzees, planned outside the context of tool-use, thus challenging the idea that planning in these species is purely domain-specific.     

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.     

Giving a Reason for Unfairness: Effects of a Rationale upon Australian Students' Performances Within an Implicit Reward Situation

Two experimental conditions (giving a rationale vs no-rationale for “unfair” rewards) were compared under an implicit reward paradigm in which “target” subjects received rewards directly and contingently for improvements on a simple motor skills task while “peer” subjects who were also performing the same task received no rewards. Data showed that there were significant reinforcement effects on the behavior of both target and peer subjects during the implicit reward situation when no rationale was given for the “unfair” application of rewards. However, when a rationale was given for the non-reward of the peer subjects, neither target nor peer subjects showed significant increases in responses from baseline. Implications for group reinforcement practices are discussed.     

Chimpanzees (Pan troglodytes) transfer tokens repeatedly with a partner to accumulate rewards in a self-control task

There has been extensive research investigating self-control in humans and nonhuman animals, yet we know surprisingly little about how one’s social environment influences self-control. The present study examined the self-control of chimpanzees in a task that required active engagement with conspecifics. The task consisted of transferring a token back and forth with a partner animal in order to accumulate food rewards, one item per token transfer. Self-control was required because at any point in the trial, either chimpanzee could obtain their accumulated rewards, but doing so discontinued the food accumulation and ended the trial for both individuals. Chimpanzees readily engaged the task and accumulated the majority of available rewards before ending each trial, and they did so across a number of conditions that varied the identity of the partner, the presence/absence of the experimenter, and the means by which they could obtain rewards. A second experiment examined chimpanzees’ self-control when given the choice between immediately available food items and a potentially larger amount of rewards that could be obtained by engaging the token transfer task with a partner. Chimpanzees were flexible in their decision-making in this test, typically choosing the option representing the largest amount of food, even if it involved delayed accumulation of the rewards via the token transfer task. These results demonstrate that chimpanzees can exhibit self-control in situations involving social interactions, and they encourage further research into this important aspect of the self-control scenario.     

Selfish strategies develop in social problem situations in chimpanzee (Pan troglodytes) mother–infant pairs

Humans employ various strategies, including selfish and altruistic strategies, depending on the situation. In order to examine whether non-human animals show such flexibility or not, we analyzed chimpanzees’ selfish and cooperative behavior in two types of social problem situations. In this study, we tested chimpanzee mother–infant pairs in two adjacent booths, each equipped with a vending machine. When a token was inserted into a vending machine, the vending machine delivered food rewards to the adjacent booth. In experiment 1, a partition between the two booths was open. In experiment 2, the partition was closed and a mother and her infant were placed in separate booths, so that reciprocal cooperation was essential for them to receive rewards. The participants did not cooperate reciprocally in either experiment. In experiment 1, the chimpanzees developed selfish tactics to get rewards and changed their tactics flexibly according to the partner’s behaviors. In experiment 2, in which they could not receive rewards without cooperation, they stopped altogether inserting tokens. In both cases, the infants stopped cooperating first. These findings support the idea that chimpanzees are primarily competitive rather than cooperative. Chimpanzees’ high social intelligence might be demonstrated in the flexibility of their selfish tactics, but not in the form of reciprocal cooperation at least when food is involved. We suggest that the failure to establish reciprocal cooperation was due to the social relationship between the mother and her infant, which was characterized by infant’s privilege and mother’s tolerance.     

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.     

Monkeys fail to reciprocate in an exchange task

Exchanges form the basis of human economies. Animals too can engage in reciprocal interactions but they do not barter goods like humans, which raises the question of the abilities necessary for trading to occur. Previous studies have shown that non-human primates can exchange food with human partners. Here, we tested the ability of brown capuchin monkeys and Tonkean macaques to reciprocate in a task requiring two conspecifics to exchange tokens in order to obtain rewards from an experimenter. We recorded 56 transfers between subjects in capuchin monkeys and 10 in Tonkean macaques. All transfers were passive in both species. Capuchins preferentially picked up tokens valuable for them in the partner’s compartment. They tended to manipulate the partner-valued tokens more often than the no-value ones, leading to more opportunities for these tokens to end up within reach of the partner. Despite optimal conditions where values of goods were defined and known by partners, however, none of the pairs tested engaged in short-term reciprocal interactions. These results indicate that calculated reciprocity was difficult if not impossible in the animals tested.     

Prospective memory in children and chimpanzees

Prospective memory (PM) involves remembering to do something at a specific time in the future. Here, we investigate the beginnings of this ability in young children (3-year-olds; Homo sapiens) and chimpanzees (Pan troglodytes) using an analogous task. Subjects were given a choice between two toys (children) or two food items (chimpanzees). The selected item was delivered immediately, whereas the unselected item was hidden in an opaque container. After completing an ongoing quantity discrimination task, subjects could request the hidden item by asking for it (children) or by pointing to the container and identifying the item on a symbol board (chimpanzees). Children and chimpanzees showed evidence of prospective-like memory in this task, as evidenced by successful retrieval of the item at the end of the task, sometimes spontaneously with no prompting from the experimenter. These findings contribute to our understanding of PM from an ontogenetic and comparative perspective.     

Are monkeys able to plan for future exchange?

Whether or not non-human animals can plan for the future is a hotly debated issue. We investigate this question further and use a planning-to-exchange task to study future planning in the cooperative domain in two species of monkeys: the brown capuchin (Cebus apella) and the Tonkean macaque (Macaca tonkeana). The rationale required subjects to plan for a future opportunity to exchange tokens for food by collecting tokens several minutes in advance. Subjects who successfully planned for the exchange task were expected to select suitable tokens during a collection period (5/10?min), save them for a fixed period of time (20/30?min), then take them into an adjacent compartment and exchange them for food with an experimenter. Monkeys mostly failed to transport tokens when entering the testing compartment; hence, they do not seem able to plan for a future exchange with a human partner. Three subjects did however manage to solve the task several times, albeit at very low rates. They brought the correct version of three possible token types, but rarely transported more than one suitable token at a time. Given that the frequency of token manipulation predicted transport, success might have occurred by chance. This was not the case, however, since in most cases subjects were not already holding the token in their hands before they entered the testing compartment. Instead, these results may reflect subjects' strengths and weaknesses in their time-related comprehension of the task.     

Precurrent self-prompting operants in children: "Remembering"

In Experiment I, one of three forms of collateral behavior was trained: Differential collateral behavior specific in form to one of two discriminative stimuli; Common collateral behavior of a single form regardless of the stimulus; or Nondifferential collateral behavior of either form regardless of the stimulus. Children were next given a short-delay matching-to-sample task in which the discriminative stimuli served as samples, and the children's previously trained collateral behavior terminated the delay and presented the comparison stimuli. Subjects engaging in sample-specific collateral behavior immediately acquired matching. Subjects engaging in sample-nonspecific collateral behavior failed to acquire matching or did so gradually. In Experiment II the minimal delay in the matching task was varied in a mixed sequence, first with collateral behavior required, and then with collateral behavior prohibited. When emitting collateral behavior Common and Nondifferential subjects showed delay-related decrements in matching while Differential subjects did not. When not emitting collateral behavior all subjects showed delay-related decrements in matching. Common and Nondifferential subjects matched more accurately when prohibited from emitting collateral behavior. Differential subjects matched more accurately when emitting collateral behavior. The results accord with Skinner's (1953, 1968) analysis of precurrent operants.     

Waiting by mistake: Symbolic representation of rewards modulates intertemporal choice in capuchin monkeys,preschool children and adult humans

In the Delay choice task subjects choose between a smaller immediate option and a larger delayed option. This paradigm, also known as intertemporal choice task, is frequently used to assess delay tolerance, interpreting a preference for the larger delayed option as willingness to wait. However, in the Delay choice task subjects face a dilemma between two preferred responses: “go for more” (i.e., selecting the larger, but delayed, option) vs. “go for sooner” (i.e., selecting the immediate, but smaller, option). When the options consist of visible food amounts, at least some of the choices of the larger delayed option might be due to a failure to inhibit a prepotent response towards the larger option rather than to a sustained delay tolerance. To disentangle this issue, we tested 10 capuchin monkeys, 101 preschool children, and 88 adult humans in a Delay choice task with food, low-symbolic tokens (objects that can be exchanged with food and have a one-to-one correspondence with food items), and high-symbolic tokens (objects that can be exchanged with food and have a one-to-many correspondence with food items). This allows evaluating how different methods of representing rewards modulate the relative contribution of the “go for more” and “go for sooner” responses. Consistently with the idea that choices for the delayed option are sometimes due to a failure at inhibiting the prepotent response for the larger quantity, we expected high-symbolic tokens to decrease the salience of the larger option, thus reducing “go for more” responses. In fact, previous findings have shown that inhibiting prepotent responses for quantity is easier when the problem is framed in a symbolic context. Overall, opting for the larger delayed option in the visible-food version of the Delay choice task seems to partially result from an impulsive preference for quantity, rather than from a sustained delay tolerance. In capuchins and children high-symbolic stimuli decreased the individual’s preference for the larger reward by distancing from its appetitive features. Conversely, the sophisticated symbolic skills of adult humans prevented the distancing effect of high-symbolic stimuli in this population, although this result may be due to methodological differences between adult humans and the other two populations under study. Our data extend the knowledge concerning the influence of symbols on both human and non-human primate behavior and add a new element to the interpretation of the Delay choice task. Since high-symbolic stimuli decrease the individual’s preference for the larger reward by eliminating those choices due to prepotent responses towards the larger quantity, they allow to better discriminate responses based on genuine delay aversion. Thus, these findings invite greater caution in interpreting the results obtained with the visible-food version of the Delay choice task, which may overestimate delay tolerance.     

Chimpanzees (Pan troglodytes) can wait,when they choose to: a study with the hybrid delay task

Self-control has been studied in nonhuman animals using a variety of tasks. The inter-temporal choice (ITC) task presents choices between smaller–sooner (SS) and larger–later (LL) options. Using food amounts as rewards, this presents two problems: (a) choices of the LL option could either reflect self-control or instead result from animals’ difficulty with pointing to smaller amounts of food; (b) there is no way to verify whether the subjects would not revert their choice for the LL option, if given the opportunity to do so during the ensuing delay. To address these problems, we have recently introduced a new protocol, the hybrid delay task, which combines an initial ITC with a subsequent accumulation phase in which selection of the SS option leads to its immediate delivery, but choice of the LL option then leads to one-by-one presentation of those items that continues only as long as the subject does not eat any of the accumulated items. The choice of the LL option therefore only reflects self-control when the number of items obtained from LL choices during the accumulation phase is higher than what could be received in the SS option. Previous research with capuchin monkeys demonstrated that their apparent self-control responses in the ITC task may have overestimated their general self-control abilities, given their poor performance in the hybrid delay task. Here, chimpanzees instead demonstrated that their choices for the LL option in the ITC phase of the hybrid delay task were confirmed by their ability to sustain long delays during accumulation of LL rewards.     

Do chimpanzees tailor their gestural signals to fit the attentional states of others?

The use of vocalizations and tactile gestures by seven juvenile chimpanzees was experimentally investigated. The subjects interacted with an experimenter who typically handed them food rewards. In some trials, however, the experimenter waited 20 s before doing so. In these trials the experimenter’s eyes were either open or closed, or the experimenter was either looking away from the subject or looking directly at him/her inquisitively with head movements. Although the chimpanzees produced at least one of the non-visual gestures mentioned (touching/tapping the experimenter or vocalizing) in 72% of all experimental trials, these actions and vocalizations were deployed without regard to the attentional state of their potential recipient, despite evidence that the subjects noticed the postures that defined the experimenter’s attentional state. The results are discussed in the context of the distinction between the evolution of an understanding of seeing/attention as an internal mental state versus an understanding of behavioral postures alone. Recieved: 12 February 1999 / Accepted after revision: 18 August 1999     

Perception of complex geometric figures in chimpanzees (Pan troglodytes) and humans (Homo sapiens): analyses of visual similarity on the basis of choice reaction time.

In a conditional-discrimination task (matching-to-sample), we assessed similarities among figures consisting of 2 elemental figures through the choice reaction time, nonmetric multidimensional scaling, and hierarchical cluster analysis data from chimpanzees (Pan troglodytes) and humans (Homo sapiens). Humans also rated similarities among figures. The results of the 3 experiments clearly indicated that the reaction time data obtained from chimpanzees' performances were useful measures of the similarities among figures. The results suggested that chimpanzees and humans perceived the complex figures similarly. The outer-contour elements were perceived most dominantly by both species, and the straight-line elements were perceived least dominantly. Both species showed the same perceptual hierarchy or dominance among perceptual categories, as determined by the similarity of simple elements, on the basis of transformational invariances.     

Scaling Incentives: Determining Preference and Indifference in 8- to lO-Year-Old Girls

The present study adopted procedures similar to those used by Logan (1965) to determine choice behavior in children. The objective was to provide an improved methodology in delay-of-gratification studies, thus avoiding the problems associated with scaling different kinds of rewards, and to provide approximate functions relating a delay-of-token reward to choice behavior. Thus, 7 girls aged 8 to 10 years were exposed to a choice paradigm in which a larger reward (2 tokens) was pitted against a smaller reward (1 token); access to these rewards was delayed a certain period of time. The results of this experiment showed that as the delay of the larger reward increased, preference for the smaller reward increased in an orderly fashion. The finding that delay shifted preference from the larger to the smaller reward is discussed in relation to current theory. The results of this experiment also provide evidence for the utility of tokens in scaling incentives for choice studies.     

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.     

Contrafreeloading and the value of control over visual stimuli in Japanese macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

Contrafreeloading, which means that animals work for food even though identical food is freely available, has been reported in animals’ feeding behavior. This phenomenon has been assumed to be explained by the information primacy model, in which the information about a food resource as well as the food itself is valuable for animals. This study confirmed a contrafreeloading-like phenomenon using movies as rewards rather than food in Japanese macaques (Macaca fuscata) and investigated the motivational system that exists behind contrafreeloading. In the experiment, movies that were presented dependently on subjects’ responses (earned movies) and movies that were presented automatically (free movies) were supplied simultaneously. The subjects continued to make responses to obtain the presentation of the earned movies although identical movies were available as free movies. These results provide the first evidence of contrafreeloading that occurs with movie rewards. The motivation maintaining the contrafreeloading behavior for movies may be control over the environment according to the competence theory.