How prosimian primates represent tools: experiments with two lemur species (Eulemur fulvus and Lemur catta)

The authors examined how 2 lemur species (Eulemur fulvus and Lemur catta) reason about tools. Experiment 1 allowed subjects to use 1 of 2 canes to retrieve an inaccessible food reward. Lemurs learned to solve this problem as quickly as other primates. Experiment 2 then presented subjects with novel tools differing from the originals along one featural dimension. Subjects attended more to tools' sizes than to their colors and made no distinction between tools' shapes and textures. Experiments 3 and 4 presented problems in which some of the tools' orientations had to be modified relative to the food. Subjects performed well on these problems, sometimes modifying the position of the tool. These results are discussed in light of the performance of other primates on this task.     

Tool choice on the basis of rigidity in capuchin monkeys

Wild capuchin monkeys select stone tools to crack open nuts on the basis of their weight and friability, two non-visual functional properties. Here, we investigated whether they would select new stick-like tools on the basis of their rigidity. In Experiment 1, subjects faced an out-of-reach reward and a choice of three unfamiliar tools differing in color, diameter, material, and rigidity. In order to retrieve the reward, capuchins needed to select the rigid tool exemplar. Capuchins gathered information regarding tools’ pliability either by (1) manipulating the tools themselves (manipulation condition), (2) observing a human demonstrator repeatedly bending the tools (observation condition), or (3) seeing the tools placed on a platform without any manipulation taking place (visual static condition). Subjects selected the rigid tool above chance levels in both the manipulation and observation conditions, but not in the visual static condition. In Experiment 2, subjects needed to select and use a flexible tool to access a liquid reward (as opposed to the rigid tool, as in previous experiment). Again, capuchins selected above chance levels the appropriate tool (i.e., flexible), thus demonstrating a good appreciation of the relation between the tool properties and the task requirements.     

Transport of tools to food sites in tufted capuchin monkeys (<Emphasis Type="Italic">Cebus apella</Emphasis>)

Tool use and transport represent cognitively important aspects of early hominid evolution, and nonhuman primates are often used as models to examine the cognitive, ecological, morphological and social correlates of these behaviors in order to gain insights into the behavior of our early human ancestors. In 2001, Jalles-Filho et al. found that free-ranging capuchin monkeys failed to transport tools (stones) to food sites (nuts), but transported the foods to the tool sites. This result cast doubt on the usefulness of Cebus to model early human tool-using behavior. In this study, we examined the performance of six captive tufted capuchin monkeys (Cebus apella) in a tool transport task. Subjects were provided with the opportunity to transport two different tools to fixed food reward sites when the food reward was visible from the tool site and when the food reward was not visible from the tool site. We found that the subjects quickly and readily transported probing tools to an apparatus baited with syrup, but rarely transported stones to a nut-cracking apparatus. We suggest that the performance of the capuchins here reflects an efficient foraging strategy, in terms of energy return, among wild Cebus monkeys.     

Tool-use learning in Tonkean macaques (Macaca tonkeana)

The transmission of tool use is a rare event in monkeys. Such an event arose in a group of semi-free-ranging Tonkean macaques (Macaca tonkeana) in which leaning a pole against the parks fence (branch leaning) appeared and spread to several males. This prompted us to test individual and social learning of this behavior in seven young males. In the first experiment, three males learned individually to obtain a food reward using a wooden pole as a climbing tool. They began using the pole to retrieve the reward only when they could alternatively experience acting on the object and reaching the target. In a second experiment, we first tested whether four other subjects could learn branch leaning after having observed a group-mate performing the task. Despite repeated opportunities to observe the demonstrator, they did not learn to use the pole as a tool. Hence we exposed the latter subjects to individual learning trials and they succeeded in the task. Tool use was not transmitted in the experimental situation, which contrasts with observations in the park. We can conclude that the subjects were not able to recognize the target as such. It is possible that they recognized it and learned the task individually when we alternated the opportunity to act upon the object and to reach the reward. This suggests that these macaques could then have associated the action they exercised upon the pole and the use of the pole as a means to reach the reward.     

Orangutans (Pongo spp.) may prefer tools with rigid properties to flimsy tools

Preference for tools with either rigid or flexible properties was explored in orangutans (Pongo spp.) through an extension of D. J. Povinelli, J. E. Reaux, and L. A. Theall's (2000) flimsy-tool problem. Three captive orangutans were presented with three unfamiliar pairs of tools to solve a novel problem. Although each orangutan has spontaneously used tools in the past, the tools presented in this study were novel to the apes. Each pair of tools contained one tool with rigid properties (functional) and one tool with flimsy properties (nonfunctional). Solving the problem required selection of a rigid tool to retrieve a food reward. The functional tool was selected in nearly all trials. Moreover, two of the orangutans demonstrated this within the first test trials with each of the three tool types. Although further research is required to test this statistically, it suggests either a preexisting preference for rigid tools or comprehension of the relevant features required in a tool to solve the task. The results of this study demonstrate that orangutans can recognize, or learn to recognize, relevant tool properties and can choose an appropriate tool to solve a problem.     

Tool selectivity in a non-primate,the New Caledonian crow (Corvus moneduloides)

We present an experiment showing that New Caledonian crows are able to choose tools of the appropriate size for a novel task, without trial-and-error learning. This species is almost unique amongst all animal species (together with a few primates) in the degree of use and manufacture of polymorphic tools in the wild. However, until now, the flexibility of their tool use has not been tested. Flexibility, including the ability to select an appropriate tool for a task, is considered to be a hallmark of complex cognitive adaptations for tool use. In experiment 1, we tested the ability of two captive birds (one male, one female), to select a stick (from a range of lengths provided) matching the distance to food placed in a horizontal transparent pipe. Both birds chose tools matching the distance to their target significantly more often than would be expected by chance. In experiment 2, we used a similar task, but with the tools placed out of sight of the food pipe, such that the birds had to remember the distance of the food before selecting a tool. The task was completed only by the male, who chose a tool of sufficient length significantly more often than chance but did not show a preference for a matching length. Electronic Publication     

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

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

Means-means-end tool choice in cotton-top tamarins (Saguinus oedipus): finding the limits on primates’ knowledge of tools

Most studies of animal tool use require subjects to use one object to gain access to a food reward. In many real world situations, however, animals perform more than one action in sequence to achieve their goals. Of theoretical interest is whether animals have the cognitive capacity to recognize the relationship between consecutive action sequences in which there may be one overall goal and several subgoals. Here we ask if cotton-top tamarins, a species that in captivity uses tools to solve means-end problems, can go one step further and use a sequence of tools (means) to obtain food (end). We first trained subjects to use a pulling tool to obtain a food reward. After this initial training, subjects were presented with problems in which one tool had to be used in combination with a second in order to obtain food. Subjects showed great difficulty when two tools were required to obtain the food reward. Although subjects attended to the connection between the tool and food reward, they ignored the physical connection between the two tools. After training on a two-tool problem, we presented subjects with a series of transfer tests to explore if they would generalize to new types of connections between the tools. Subjects readily transferred to new connections. Our results therefore provide the first evidence to date that tamarins can learn to solve problems involving two tools, but that they do so only with sufficient training.     

Donor payoffs and other-regarding preferences in cotton-top tamarins (Saguinus oedipus)

Helping others at no cost to oneself is a simple way to demonstrate other-regarding preferences. Yet, primates exhibit mixed results for other-regarding preferences: chimpanzees and tamarins do not show these effects, whereas capuchin monkeys and marmosets preferentially give food to others. One factor of relevance to this no-cost food donation is the payoff to the donor. Though donors always receive the same payoffs regardless of their choice, previous work varies in whether they receive either a food reward or no food reward. Here, I tested cotton-top tamarins in a preferential giving task. Subjects could choose from two tools, one of which delivered food to a partner in an adjacent cage and the other of which delivered food to an empty cage. Thus, subjects could preferentially give or withhold food from a partner. I varied whether subjects received food payoffs, whether a partner was present or absent, and whether the partner was a non-cagemate or the subject’s mate. Results showed that the subjects’ overall motivation to pull either tool declined when they did not receive any food. Additionally, they did not preferentially donate or withhold food, regardless of their own payoff or their relationship with the partner. Thus, cotton-top tamarins do not take advantage of cost-free food giving, either when they might gain in the future (mates) or when they have no opportunity for future interactions (non-cagemates).     

Dogs are able to solve a means-end task

Dogs, although very skilled in social-communicative tasks, have shown limited abilities in the domain of physical cognition. Consequently, several researchers hypothesized that domestication enhanced dogs’ cognitive abilities in the social realm, but relaxed selection on the physical one. For instance, dogs failed to demonstrate means-end understanding, an important form of relying on physical causal connection, when tested in a string-pulling task. Here, we tested dogs in an “on/off” task using a novel approach. Thirty-two dogs were confronted with four different conditions in which they could choose between two boards one with a reward “on” and another one with a reward “off” (reward was placed next to the board). The dogs chose the correct board when (1) both rewards were placed at the same distance from the dog, when (2) the reward placed “on” the board was closer to the dog, and (3) even when the reward placed “off” the board was much closer to the dog and was food. Interestingly, in the latter case, dogs did not perform above chance, if instead of a direct reward, the dogs had to retrieve an object placed on the board to get a food reward. In contrast to previous string-pulling studies, our results show that dogs are able to solve a means-end task even if proximity of the unsupported reward is a confounding factor.     

Causal knowledge and imitation/emulation switching in chimpanzees (Pan troglodytes) and children (Homo sapiens)

This study explored whether the tendency of chimpanzees and children to use emulation or imitation to solve a tool-using task was a response to the availability of causal information. Young wild-born chimpanzees from an African sanctuary and 3- to 4-year-old children observed a human demonstrator use a tool to retrieve a reward from a puzzle-box. The demonstration involved both causally relevant and irrelevant actions, and the box was presented in each of two conditions: opaque and clear. In the opaque condition, causal information about the effect of the tool inside the box was not available, and hence it was impossible to differentiate between the relevant and irrelevant parts of the demonstration. However, in the clear condition causal information was available, and subjects could potentially determine which actions were necessary. When chimpanzees were presented with the opaque box, they reproduced both the relevant and irrelevant actions, thus imitating the overall structure of the task. When the box was presented in the clear condition they instead ignored the irrelevant actions in favour of a more efficient, emulative technique. These results suggest that emulation is the favoured strategy of chimpanzees when sufficient causal information is available. However, if such information is not available, chimpanzees are prone to employ a more comprehensive copy of an observed action. In contrast to the chimpanzees, children employed imitation to solve the task in both conditions, at the expense of efficiency. We suggest that the difference in performance of chimpanzees and children may be due to a greater susceptibility of children to cultural conventions, perhaps combined with a differential focus on the results, actions and goals of the demonstrator.     

Probing the limits of tool competence: Experiments with two non-tool-using species (Cercopithecus aethiops and Saguinus oedipus)

Non-human animals vary in their ability to make and use tools. The goal of the present study was to further explore what, if anything, differs between tool-users and non-tool-users, and whether these differences lie in the conceptual or motor domain. We tested two species that typically do not use tools-cotton top tamarins (Saguinus oedipus) and vervet monkeys (Cercopithecus aethiops)-on problems that mirrored those designed for prolific tool users such as chimpanzees. We trained subjects on a task in which they could choose one of two canes to obtain an out-of-reach food reward. After training, subjects received several variations on the original task, each designed to examine a specific conceptual aspect of the pulling problem previously studied in other tool-using species. Both species recognized that effective pulling tools must be made of rigid materials. Subsequent conditions revealed significant species differences, with vervets outperforming tamarins across many conditions. Vervets, but not tamarins, had some recognition of the relationship between a tool's orientation and the position of the food reward, the relationship between a tool's trajectory and the substance that it moves on, and that tools must be connected in order to work properly. These results provide further evidence that tool-use may derive from domain-general, rather than domain-specific cognitive capacities that evolved for tool use per se.     

Big brains are not enough: performance of three parrot species in the trap-tube paradigm

The trap-tube task has become a benchmark test for investigating physical causality in vertebrates. In this task, subjects have to retrieve food out of a horizontal tube using a tool and avoiding a trap hole in the tube. Great apes and corvids succeeded in this task. Parrots with relative brain volumes comparable to those of corvids and primates also demonstrate high cognitive abilities. We therefore tested macaws, a cockatoo, and keas on the trap-tube paradigm. All nine parrots failed to solve the task. In a simplified task, trap tubes with a slot inserted along the top were offered. The slot allowed the birds to move the reward directly with their bills. All but one individual solved this task by lifting the food over the trap. However, the parrots failed again when they were prevented from lifting the reward, although they anticipated that food will be lost when moved into the trap. We do not think that the demanding use of an external object is the main reason for the parrots’ failure. Moreover, we suppose these parrots fail to consider the trap’s position in the beginning of a trial and were not able to stop their behaviour and move the reward in the trap’s opposite direction.     

Juvenile domestic pigs (Sus scrofa domestica) use human-given cues in an object choice task

Research on the comprehension of human-given cues by domesticated as well as non-domesticated species has received considerable attention over the last decade. While several species seem to be capable of utilizing these cues, former work with domestic pigs (Sus scrofa domestica) has shown inconclusive results. In this study, we investigated the use of human-given cues in an object choice task by young domestic pigs (N = 17; 7 weeks of age) who had very limited human contact prior to the experiments. Subjects had to choose between two bowls of which only one was baited with a reward. Over the course of five experiments, pigs were able to use proximal and, with some constraints, also distal pointing cues presented in both a dynamic-sustained and in a momentary manner. When the experimenter was pointing from the incorrect bowl towards the correct one, most of the subjects had problems solving the task—indicating that some form of stimulus/local enhancement affected pigs’ decision making. Interestingly, pigs were able to utilize the body and head orientation of a human experimenter to locate the hidden reward but failed to co-orient when head or body orientation of the experimenter was directed into distant space with no bowls present. Control trials ruled out the possibility that other factors (e.g. odour cues) affected subjects’ choice behaviour. Learning during experiments played a minor role and only occurred in three out of twelve test conditions. We conclude that domestic pigs, even at a very young age, are skilful in utilizing various human-given cues in an object choice task—raising the question whether pigs only used stimulus/local enhancement and associative learning processes or whether they were able to comprehend the communicative nature of at least some of these cues.     

The role of the GABA(A) agonist muscimol on memory performance: reward contingencies determine the nature of the deficit

A matching-to-position (MTP) paradigm was altered to influence the type of associations a rat would use to solve the task. Our main behavioral manipulation was the application of the differential outcomes procedure (DOP). The DOP involves correlating each to-be-remembered event with a distinct reward condition. This procedure results in the development of unique reward expectancies that enhance and guide choice behavior. Such distinct reward expectancies are not formed when either a common or random assignment of reward is used (a non-differential outcomes procedure [NOP]). Intracerebroventricular infusions of the amnestic agent muscimol (GABA(A) agonist) or aCSF were delivered to male rats trained on a delayed MTP task that implemented either the DOP or the NOP. Muscimol impaired performance in a dose dependent fashion in both groups--but the nature of the deficit differed as a function of reinforcement contingencies. Rats trained with the DOP displayed a non-mnemonic delay-independent impairment: performance at all delay intervals was disrupted. In contrast, NOP-trained rats displayed a delay-dependent impairment demonstrating that muscimol can also have memory-disrupting effects. The difference in pattern of impairment appears to be a function of the associations formed during training and the type of cognitive strategies involved in maintaining behavior on a conditional delayed discrimination task when reinforcement contingencies are varied. Thus, these results demonstrate that increasing GABA(A) receptor activation impairs a range of associative and memory functions.     

Interpersonal and intrapersonal justice: The effect of subject and confederate outcomes on evaluations of fairness

Subjects were asked to make justice judgments based on different comparison standards—intrapersonal and interpersonal standards. The intrapersonal standard of entitlement was induced through a promise. Subjects were either given more than ($4.00), less than ($1.00), or exactly ($2.50) the reward promised for completion of an assigned task. By providing subjects access to information pertaining to a confederate co-worker's outcomes, subjects were able to evaluate the equitableness (an interpersonal standard) of the distribution. Confederates also received one of three levels of reward (i.e. $1.00, $2.50, or $4.00). It was found that evaluations of fairness depend on the criteria used in making the assessment; if intrapersonal comparisons are employed, subjects produce justice judgments quite different from those grounded on an interpersonal referent. A main effect for subject outcome was obtained when the prevailing standard was intrapersonal and an interaction between subject and confederate outcomes was found when the prevailing standard was interpersonal. One implication of these results is that it is possible for an equitable distribution to still be considered unjust.     

Understanding the functional properties of tools: chimpanzees (Pan troglodytes) and capuchin monkeys (Cebus apella) attend to tool features differently

We examined whether eight capuchins and eight chimpanzees were able to retrieve a reward placed inside a tube, of varying length, by selecting the correct stick from different sets of three sticks differing in length (functional feature) and handle (non-functional feature). Moreover, to investigate whether seeing the stick inside the tube (visual feedback) improves performance, half of the subjects were tested with a transparent apparatus and the other half with an opaque apparatus. Phase 1 included (a) Training 1 in which each stick had a different handle and (b) Transfer 1 in which the handles were switched among sticks, so that the functional tool had the same length but a different handle than before. The seven chimpanzees and one capuchin that passed Transfer 1 received Transfer 2. The other subjects received (a) Training 2, which used the same sticks from Phase 1 with handles switched in every trial, and (b) Transfer 2 in which the tube was longer, all sticks had the same new handle, and the formerly longest tool became intermediate in length. Eight chimpanzees and three capuchins passed Transfer 2. Results showed that (1) chimpanzees applied relational structures in tool using tasks more quickly than capuchins and (2) capuchins required more varied experience to attend to the functional feature of the tool. Interestingly, visual feedback did not improve performance in either species.     

Undermining the Zeigarnik effect: Another hidden cost of reward

Forty college students participated in a study ostensibly designed to obtain measures of the hemispheric activity while they worked on a spatial reasoning task. In fact, our true interest was in determining whether subjects would return to the spatial reasoning task once the hemispheric recordings were completed. Subjects would normally be expected to return to the task by virtue of the Zeigarnik effect because no subject completed the task during the “hemispheric recording” phase. The manipulation involved telling one group of subjects that they would be paid $1.50 for participating in the study (expected reward group). The remainder of the subjects were not led to expect the reward (unexpected reward group). The result was that 86% of the unexpected reward subjects but only 58% of the expected reward subjects (p < .05) manifested the Zeigarnik effect. This differential tendency to return to the task was further reflected in time differences. The average unexpected reward subject spent 3 min 48 sec of a five-minute free-choice period at the task, whereas the average expected reward subject spent only 2 min 20 sec (p < .05). Reward expectancy, therefore, led to an undermining of the Zeigarnik effect. This observation supports Condry's prediction that rewarding performance at a task can lead to premature task disengagement.     

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.     

Observer states and traits and the imitative process: A test of a new formulation

In three studies the formulation was tested that characteristics of the observer have an effect on imitation only in situations which provide little information concerning appropriate or expected behaviors. In Experiment I, female college students of high, medium, and low need for social approval underwent either a high or low arousal manipulation. Subjects observed a videotaped model who engaged in a size judgment task. For half the subjects the model received verbal reward (high information); for the other half the model received no feedback (low information). Subsequently, the model and subject responded alternately in the task during which no feedback was given. Neither need for social approval nor manipulated arousal was found to be systematically related to imitation in either information condition. However, the hypothesized relationships were found when subjects' arousal or anxiety levels were determined by self ratings. These results were replicated in a second, similar experiment. In a third experiment, which involved an improved arousal manipulation, the hypothesized relationships were confirmed.