The relationship between problem solving and inhibitory control: cotton-top tamarin (Saguinus oedipus) performance on a reversed contingency task

To explore the relationship between problem solving and inhibitory control, the authors present 4 experiments on cotton-top tamarins (Saguinus oedipus) using a reverse-reward contingency task. In Experiment 1, 1 group of tamarins was given a choice between a small and a large quantity of food. Whichever quantity the tamarins reached for first, they received the alternative. The tamarins consistently picked the larger quantity, thereby receiving the smaller. A 2nd group of tamarins was given the same task, except that if they reached for the larger quantity of food, they received nothing. The tamarins continued to pick the larger quantity, even though this resulted in no food. In addition, most of the tamarins continued to pick the larger quantity even when the food payoff for choosing the smaller quantity was increased (Experiment 2) or when the visual salience of the food was reduced (Experiment 3). Experiment 4 was based on the finding that chimpanzees (Pan troglodytes) that have been trained on the concept of number can solve the reversed contingency task if the food is replaced by Arabic numerals. With the help of a color association, and a higher cost incurred by picking the color associated with 3 food items, the tamarins learned to pick the color associated with 1 food item. These results are compared with those obtained from studies of other primate species, highlighting the importance of comparative studies of problem solving that use comparable methods.     

Behavioral contrast in competitive and noncompetitive environments

Three experiments examined the effects of opportunities for an alternative response (drinking) on positive behavioral contrast of rats' food-reinforced bar pressing. In both Experiments 1 and 2 the baseline multiple variable-interval schedules were rich (variable interval 10-s), and contrast was examined both with and without a water bottle present. In Experiment 1, the rats were not water deprived. When one component of the multiple schedule was changed to extinction, the rate of bar pressing increased in the constant component (positive behavioral contrast). The magnitude of contrast was larger when the bottle was absent than when it was present, as predicted by the matching law. Drinking did not shift from the constant variable-interval component to the extinction component, as might have been expected from competition theory. In Experiment 2, the rats were water deprived. Contrast was larger when the bottle was present than when it was absent, and drinking did shift to the extinction component, as predicted by competition theory. In Experiment 3, water-deprived rats responded on leaner multiple variable-interval schedules (60-s) in the presence of a water bottle. When one component was changed to extinction, contrast did not occur, and drinking did not shift to the extinction component. The present results suggest that there are at least two different sources of behavioral contrast: “competitive” contrast, observed when an alternative response occurs with high probability, and “noncompetitive” contrast, observed when an alternative response occurs with low probability. The results, in conjunction with earlier studies, also suggest that the form of the alternative response and the rate of food reinforcement provided by the multiple schedule combine to determine the amount of contrast.     

Quantity-based judgments in the domestic dog (<Emphasis Type="Italic">Canis lupus familiaris</Emphasis>)

We examined the ability of domestic dogs to choose the larger versus smaller quantity of food in two experiments. In experiment 1, we investigated the ability of 29 dogs (results from 18 dogs were used in the data analysis) to discriminate between two quantities of food presented in eight different combinations. Choices were simultaneously presented and visually available at the time of choice. Overall, subjects chose the larger quantity more often than the smaller quantity, but they found numerically close comparisons more difficult. In experiment 2, we tested two dogs from experiment 1 under three conditions. In condition 1, we used similar methods from experiment 1 and tested the dogs multiple times on the eight combinations from experiment 1 plus one additional combination. In conditions 2 and 3, the food was visually unavailable to the subjects at the time of choice, but in condition 2, food choices were viewed simultaneously before being made visually unavailable, and in condition 3, they were viewed successively. In these last two conditions, and especially in condition 3, the dogs had to keep track of quantities mentally in order to choose optimally. Subjects still chose the larger quantity more often than the smaller quantity when the food was not simultaneously visible at the time of choice. Olfactory cues and inadvertent cuing by the experimenter were excluded as mechanisms for choosing larger quantities. The results suggest that, like apes tested on similar tasks, some dogs can form internal representations and make mental comparisons of quantity.     

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) perform on the reversed contingency task: the effects of food quantity and food visibility

S. T. Boysen and G. G. Berntson (1995) found that chimpanzees performed poorly on a reversed contingency task in which they had to point to the smaller of 2 food quantities to acquire the larger quantity. The authors compared the performance of 4 great ape species (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) on the reversed contingency task while manipulating food quantity (0-4 or 1-4) and food visibility (visible pairs or covered pairs). Results showed no systematic species differences but large individual differences. Some individuals of each species were able to solve the reversed contingency task. Both quantity and visibility of the food items had a significant effect on performance. Subjects performed better when the disparity between quantities was smaller and the quantities were not directly visible.     

Voluminous sucrose consumption in female rats: increased "nippiness" during periods of sucrose removal and possible oestrus periodicity

In a two-bottle paradigm in which water and 10% sucrose water were always available, female rats drank about 200 cc of the sugar water (about 65 g of sucrose/kg) per day for 4 wk. There were no significant decreases in consumption over this time. In Exp. 2 female rats singly housed were given two bottles containing water for 1 wk. and then a bottle containing water and a bottle containing 15% sucrose for the next week for 6 wk. When sucrose was available, the rats ate 33% more rat chow. When sucrose was removed, the rats displayed more episodes of biting a stimulus when the food cubes were being removed for daily measurements. Some females exhibited a marked 4- to 5-day periodicity in sucrose (7.5%) consumption. The persistent and voluminous consumption of sucrose water and enhanced agonistic-like behavior during periods of withdrawal suggests the presence of a robust phenomenon with potential dinical applications to the challenge of addiction.     

Foraging behaviour in domestic pigs (Sus scrofa): remembering and prioritizing food sites of different value

This experiment investigated whether domestic pigs can remember the locations of food sites of different relative value, and how a restricted retrieval choice affects their foraging behaviour. Nine juvenile female pigs were trained to relocate two food sites out of a possible eight in a spatial memory task. The two baited sites contained different amounts of food and an obstacle was added to the smaller amount to increase handling time. On each trial, a pig searched for the two baited sites (search visit). Once it had found and eaten the bait, it returned for a second (relocation) visit, in which the two same sites were baited. Baited sites were changed between trials. All subjects learnt the task. When allowed to retrieve both baits, the subjects showed no preference for retrieving a particular one first (experiment 1). When they were allowed to retrieve only one bait, a significant overall preference for retrieving the larger amount emerged across subjects (experiment 2). To test whether this preference reflected an avoidance of the obstacle with the smaller bait, 15 choice-restricted control trials were conducted. In control trials obstacles were present with both baits. Pigs continued to retrieve the larger bait, indicating they had discriminated between the two food sites on the basis of quantity or profitability and adjusted their behaviour accordingly when the relocation choice was restricted. This suggests for the first time that domestic pigs have the ability to discriminate between food sites of different relative value and to remember their respective locations.     

More or less: spontaneous quantity discrimination in the domestic cat

We examined spontaneous quantity discrimination in untrained domestic cats in three food choice experiments. In Experiment 1, we presented the cats with two different quantities of food in eight numerical combinations. Overall, the subjects chose the larger quantity more often than the smaller one, and significantly so when the ratio between the quantities was less than 0.5. In Experiment 2, we presented the cats with two pieces of food in four different size combinations. Again, subjects chose the larger piece above chance, although not in the combination where the largest item was presented. In Experiment 3, a subset of the cats was presented multiple times with two different quantities of food, which were hidden from view. In this case, the cats did not choose the larger quantity more often than the smaller one, suggesting that in the present experiments they mainly used visual cues when comparing quantities. We conclude that domestic cats are capable of spontaneously discriminating quantities when faced with different numbers or sizes of food items, and we suggest why they may not always be motivated to choose the larger quantity. In doing so, we highlight the advantages of testing spontaneous choice behavior, which is more likely to reflect animals’ everyday manner of responding than is the case when training them in order to test their absolute limits of performance which may not always coincide with their daily needs.     

Size matters: impact of item size and quantity on array choice by chimpanzees (Pan troglodytes)

The authors previously reported that chimpanzees (Pan troglodytes) showed a striking bias to select the larger of 2 candy arrays, despite a reversed reward contingency in which the animals received the smaller, nonselected array as a reward, except when Arabic numerals were used as stimuli. A perceptual or incentive-based interference occurred that was overcome by symbolic stimuli. The authors of the present study examined the impact of element size in choice arrays, using 1 to 5 large and small candies. Five test-sophisticated chimpanzees selected an array from the 2 presented during each trial. Their responses were not optimal, as animals generally selected arrays with larger total mass; thus, they received the smaller remaining array as a reward. When choice stimuli differed in size and quantity, element size was more heavily weighted, although choices reflected total candy mass. These results replicate previous findings showing chimpanzees' difficulties with quantity judgments under reverse reward contingencies and also show that individual item size exerts a more powerful interference effect.     

Transfer of self-control in black (Eulemur macaco) and brown (Eulemur fulvus) lemurs: choice of a less preferred food item under a reverse-reward contingency

When presented a choice between two food-type arrays of equivalent size under a reverse-reward contingency, black (Eulemur macaco) and brown (Eulemur fulvus) lemurs transposed their self-control abilities, acquired in a previous experiment, to significantly select the less-desired food item in order to gain access to the more desired one. However, when presented with the choice between two different food-type arrays in which the amount of the less desired food array was larger than the more desired one, large individual differences were revealed: Some subjects established a consistent rule favoring quality or quantity, whereas others exhibited various point of trade-off. These results show that lemurs seem to manage the task considering not only food quantity but also food quality.     

Preference reversals with food and water reinforcers in rats

Rats were given a choice between a smaller, immediately available reward and a larger reward available after a delay. In one phase, the reward was food and in another phase, the reward was water. Constant delays were added between the choice presentation and the delivery of the reward alternatives. As the time between choice and reward delivery increased from 0 to 25 s, all rats (except one in the water phase) reversed their preference from the smaller, sooner alternative to the larger, later alternative. These findings extend the generality of the preference-reversal animal model to qualitatively different reinforcers. Furthermore, the presence of both impulsive and self-control choices within the same animal is consistent with the view that self-control may be understood as choice behavior, and that species differences in self-control may be differences in degree, not kind.     

Capuchin monkeys (Cebus apella) succeed in a test of quantity conservation

Nonhuman animals demonstrate a number of impressive quantitative skills such as counting sets of items, comparing sets on the basis of the number of items or amount of material, and even responding to simple arithmetic manipulations. In this experiment, capuchin monkeys were presented with a computerized task designed to assess conservation of discrete quantity. Monkeys first were trained to select from two horizontal arrays of stimuli the one with the larger number of items. On some trials, after a correct selection there was no feedback but instead an additional manipulation of one of those arrays. In some cases, this manipulation involved moving items closer together or farther apart to change the physical arrangement of the array but not the quantity of items in the array. In other cases, additional items were added to the initially smaller array so that it became quantitatively larger. Monkeys then made a second selection from the two arrays of items. Previous research had shown that rhesus monkeys (Macaca mulatta) succeeded with this task. However, there was no condition in that study in which items were added to the smaller array without increasing its quantity to a point where it became the new larger array. This new condition was added in the present experiment. Capuchin monkeys were sensitive to all of these manipulations, changing their selections when the manipulations changed which array contained the larger number of items but not when the manipulations changed the physical arrangement of items or increased the quantity in one array without also reversing which of the two arrays had more items. Therefore, capuchin monkeys responded on the basis of the quantity of items, and they were not distracted by non-quantitative manipulations of the arrays. The data indicate that capuchins are sensitive to simply arithmetic manipulations that involve addition of items to arrays and also that they can conserve quantity.     

Relative quantity judgment by Asian elephants (Elephas maximus)

This study investigated whether Asian elephants can make relative quantity judgment (RQJ), a dichotomous judgment of unequal quantities ordered in magnitude. In Experiment 1, elephants were simultaneously shown two baskets with differing quantities of bait (up to 6 items). In Experiment 2, elephants were sequentially presented with baits, which could not be seen by elephants in their total quantities. The task of elephants was to choose the larger quantity in both experiments. Results showed that the elephants chose the larger quantity with significantly greater frequency. Interestingly, the elephants did not exhibit disparity or magnitude effects, in which performance declines with a smaller difference between quantities in a two-choice task, or the total quantity increases, respectively. These findings appear to be inconsistent with the previous reports of RQJ in other animals, suggesting that elephants may be using a different mechanism to compare and represent quantities than previously suggested for other species.     

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.     

Inhibitory control and response selection in problem solving: how cotton-top tamarins (Saguinus oedipus) overcome a bias for selecting the larger quantity of food

When presented with a choice between 1 and 3 pieces of food in a type of reversed contingency task, 4 cotton-top tamarins (Saguinus oedipus) consistently chose the 3 pieces of food and received nothing, even though the choice of 1 piece would have yielded 3. However, in a task in which the tamarins received the 1 piece of food when they chose it, all subjects learned to select 1 over 3. Thus, the tamarins' prior failure on the reversed contingency task did not result entirely from an inherent inability to suppress the prepotent response of reaching to the larger of 2 quantities of food. After the experience of selecting the smaller quantity and receiving it, all of the tamarins solved the version of the reversed contingency task that they failed initially. These results suggest that the tamarins' initial failure may have reflected a difficulty with selecting an alternative response option.     

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.     

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.     

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.     

Mangabeys (<Emphasis Type="Italic">Cercocebus torquatus lunulatus</Emphasis>) solve the reverse contingency task without a modified procedure

Problem solving often relies on generating new responses while inhibiting others, particularly prepotent ones. A paradigm to study inhibitory abilities is the reverse contingency task (Boysen and Berntson in J Exp Psychol Anim Behav Process 21:82–86, 1995), in which two different quantities of food are offered to an individual who receives the array he did not choose. Therefore, mastery of the task demands selecting the smaller quantity to obtain the larger one. Several non-human primates have been tested in the reverse contingency task. To date, only great apes and rhesus monkeys (Macaca mulatta) have succeeded in the original task, with no need of procedural modifications as the large-or-none contingency, correction trials or symbolic stimuli substituting for actual food quantities. Here, four mangabeys were presented with two stimulus arrays of one and four raisins in the context of the reverse contingency task. Three of them learned to perform the task well above chance without a modified procedure. They also reached above-chance performance when presented with two stimulus arrays of zero and four raisins, despite the initial difficulty of choosing a null quantity. After a period of 7–10 months, in which the animals were not tested on any task, all three subjects continued to perform well, even when presented with novel quantity pairs.     

Summation of response rates to discriminative stimuli associated with qualitatively different reinforcers

In Experiment I, a four-ply multiple schedule was used to study the effects on rate of responding in rats of food, water, and food and/or water reinforcement under different deprivation conditions. Food and water were associated separately with different stimuli, the combination of which was associated with food and water together, or with food or water randomly. Rates in the presence of the combined stimuli were consistently intermediate to the rates generated by the separate stimuli, a result seemingly incompatible with a "summation" hypothesis. Experiment II was a simplified systematic replication of Experiment I, verifying the major findings.     

DISCOUNTING OF DELAYED FOOD REWARDS IN PIGEONS AND RATS: IS THERE A MAGNITUDE EFFECT?

Temporal discounting refers to the decrease in the present, subjective value of a reward as the time to its receipt increases. Results from humans have shown that a hyperbola-like function describes the form of the discounting function when choices involve hypothetical monetary rewards. In addition, magnitude effects have been reported in which smaller reward amounts are discounted more steeply than larger amounts. The present research examines the cross-species generality of these findings using real rewards, namely food pellets, with both pigeons and rats. As with humans, an adjusting amount procedure was used to estimate the amount of immediate reward judged equal in value to a delayed reward. Different amounts of delayed food rewards (ranging from 5 to 32 pellets in pigeons and from 5 to 20 pellets in rats) were studied at delays varying from 1 s to 32 s. A simple hyperbola, similar to the hyperbola-like mathematical function that describes the discounting of hypothetical monetary rewards in humans, described the discounting of food rewards in both pigeons and rats. These results extend the generality of the mathematical model of discounting. Rates of discounting delayed food rewards were higher for pigeons than for rats. Unlike humans, however, neither pigeons nor rats showed a reliable magnitude effect: Rate of discounting did not vary systematically as a function of the amount of the delayed reward.