Raking it in: the impact of enculturation on chimpanzee tool use

Recent evidence for different tool kits, proposed to be based upon culture-like transmission, have been observed across different chimpanzee communities across Western Africa. In light of these findings, the reported failures by seven captive juvenile chimpanzees tested with 27 tool use tasks (Povinelli 2000) seem enigmatic. Here we report successful performance by a group of nine captive, enculturated chimpanzees, and limited success by a group of six semi-enculturated chimpanzees, on two of the Povinelli tasks, the Flimsy Tool task, and the Hybrid Tool task. All chimpanzees were presented with a rake with a flimsy head and a second rake with a rigid head, either of which could be used to attempt to retrieve a food reward that was out of reach. The rigid rake was constructed such that it had the necessary functional features to permit successful retrieval, while the flimsy rake did not. Both chimpanzee groups in the present experiment selected the functional rigid tool correctly to use during the Flimsy Tool task. All animals were then presented with two “hybrid rakes” A and B, with one half of each rake head constructed from flimsy, non-functional fabric, and the other half of the head was made of wood. Food rewards were placed in front of the rigid side of Rake A and the flimsy side of Rake B. To be successful, the chimps needed to choose the rake that had the reward in front of the rigid side of the rake head. The fully enculturated animals were successful in selecting the functional rake, while the semi-enculturated subjects chose randomly between the two hybrid tools. Compared with findings from Povinelli, whose non-enculturated animals failed both tasks, our results demonstrate that chimpanzees reared under conditions of semi-enculturation could learn to discriminate correctly the necessary tool through trial-and-error during the Flimsy Tool task, but were unable to recognize the functional relationship necessary for retrieving the reward with the “hybrid” rake. In contrast, the enculturated chimpanzees were correct in their choices during both the Flimsy Tool and the Hybrid Tool tasks. These results provide the first empirical evidence for the differential effects of enculturation on subsequent tool use capacities in captive chimpanzees.     

Discrimination of functionally appropriate and inappropriate throwing tools by captive tufted capuchins (<Emphasis Type="Italic">Cebus apella</Emphasis>)

A tool-throwing task was used to test whether capuchin monkeys understand the difference between functionally appropriate and functionally inappropriate tools. A group of monkeys was trained to obtain a sticky treat from a container outside their enclosure using a projectile attached to one end of an anchored line. Subsequently, these monkeys were given choice tests between functional and nonfunctional versions of tools used in training. A different feature of the tool was varied between alternatives in each choice test. The monkeys chose to use functional tools significantly more often than nonfunctional tools in early exposures to each choice test. A second experiment tested whether these subjects, as well as a second group of minimally trained participants, could distinguish between functional and nonfunctional tools that appeared different from those used in training. A new set of design features was varied between tools in these choice tests. All participants continued to choose functional tools significantly more often than nonfunctional tools, regardless of their tool-throwing experience or the novel appearance of the tools. These results suggest that capuchin monkeys, like chimpanzees studied in similar experiments, are sensitive to a variety of functionally relevant tool features.     

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.     

How do tufted capuchin monkeys (Cebus apella) understand causality involved in tool use?

Four tufted capuchin monkeys (Cebus apella) were trained to choose from 2 hook-like tools, 1 of which successfully led to collecting food, whereas the other did not because of inappropriate spatial arrangement of the tool and the food. In Experiment 1, all of the monkeys successfully learned the basic task. The monkeys performed successfully with tools of novel colors and shapes in Experiments 2-5. These results demonstrate that the monkeys used the spatial arrangement of the tool and the food as a cue. However, they failed when there were obstacles (Experiment 6) or traps (Experiment 7) on the path along which the monkeys dragged tools. These results may suggest that capuchin monkeys understand the spatial relationship between 2 items, namely, food and the tool, but do not understand the spatial relationship among 3 items, namely, food, tool, and the environmental condition. The possible role of stimulus generalization is also considered.     

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.     

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     

Task constraints mask great apes' ability to solve the trap-table task

Researchers have investigated animals' causal knowledge with a task requiring subjects to use a tool to bring a reward within reach whilst avoiding a trap. Previous studies have suggested limitations in the ability of several species to avoid traps in tubes or tables. However, certain features may have inflated task difficulty. We tested 20 chimpanzees (Pan troglodytes), 7 orangutans (Pongo pygmaeus), 5 bonobos (Pan paniscus), and 5 gorillas (Gorilla gorilla) in the trap-table--a task in which subjects have to pull one of two rakes prepositioned behind two rewards on a flat surface. One of the rewards is in front of a trap into which it will fall. We investigated the effect of trap type, tool type, the number of available tools, and reinforcement regime on performance. We replicated previous findings showing that apes failed to choose the correct rake above chance. However, when they could instead choose where to insert a single tool, around 80% of the apes solved the trap-table task in the first trial, revealing an important effect of task constraints on their performance.     

Do wild New Caledonian crows (<Emphasis Type="Italic">Corvus moneduloides</Emphasis>) attend to the functional properties of their tools?

New Caledonian crows are the most proficient non-hominin tool manufacturers but the cognition behind their remarkable skills remains largely unknown. Here we investigate if they attend to the functional properties of the tools that they routinely use in the wild. Pandanus tools have natural barbs along one edge that enable them to function as hooking implements when the barbs face backwards from the working tip. In experiment 1 we presented eight crows with either a non-functional (‘upside-down’) or a functional pandanus tool in a baited hole. Four of the crows never flipped the tools. The behaviour of the four flipping birds suggested that they had a strategy of flipping a tool when it was not working. Observations of two of the eight crows picking up pandanus tools at feeding tables in the wild supported the lack of attention to barb direction. In experiment 2 we gave six of the eight crows a choice of either a barbed or a barbless pandanus tool. Five of the crows chose tools at random, which further supported the findings in experiment 1 that the crows paid little or no attention to the barbs. In contrast, a third experiment found that seven out of eight crows flipped non-functional stick tools significantly more than functional ones. Our findings indicate that the crows do not consistently attend to the presence or orientation of barbs on pandanus tools. Successful pandanus tool use in the wild seems to rely on behavioural strategies formed through associative learning, including procedural knowledge about the sequence of operations required to make a successful pandanus tool. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Can orangutans (Pongo abelii) infer tool functionality?

It is debatable whether apes can reason about the unobservable properties of tools. We tested orangutans for this ability with a range of tool tasks that they could solve by using observational cues to infer tool functionality. In experiment 1, subjects successfully chose an unbroken tool over a broken one when each tool’s middle section was hidden. This prevented seeing which tool was functional but it could be inferred by noting the tools’ visible ends that were either disjointed (broken tool) or aligned (unbroken tool). We investigated whether success in experiment 1 was best explained by inferential reasoning or by having a preference per se for a hidden tool with an aligned configuration. We conducted a similar task to experiment 1 and included a functional bent tool that could be arranged to have the same disjointed configuration as the broken tool. The results suggested that subjects had a preference per se for the aligned tool by choosing it regardless of whether it was paired with the broken tool or the functional bent tool. However, further experiments with the bent tool task suggested this preference was a result of additional demands of having to attend to and remember the properties of the tools from the beginning of the task. In our last experiment, we removed these task demands and found evidence that subjects could infer the functionality of a broken tool and an unbroken tool that both looked identical at the time of choice.     

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).     

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.     

Involvement of the rat anterior cingulate cortex in control of instrumental responses guided by reward expectancy

The anterior cingulate cortex (ACC) plays a critical role in stimulus-reinforcement learning and reward-guided selection of actions. Here we conducted a series of experiments to further elucidate the role of the ACC in instrumental behavior involving effort-based decision-making and instrumental learning guided by reward-predictive stimuli. In Experiment 1, rats were trained on a cost-benefit T-maze task in which they could either choose to climb a barrier to obtain a high reward (four pellets) in one arm or a low reward (two pellets) in the other with no barrier present. In line with previous studies, our data reveal that rats with quinolinic acid lesions of the ACC selected the response involving less work and smaller reward. Experiment 2 demonstrates that breaking points of instrumental performance under a progressive ratio schedule were similar in sham-lesioned and ACC-lesioned rats. Thus, lesions of the ACC did not interfere with the effort a rat is willing to expend to obtain a specific reward in this test. In a subsequent task, we examined effort-based decision-making in a lever-press task where rats had the choice between pressing a lever to receive preferred food pellets under a progressive ratio schedule, or free feeding on a less preferred food, i.e. lab chow. Results show that sham- and ACC-lesioned animals had similar breaking points and ingested comparable amounts of less-preferred food. Together, the results of Experiment 1 and 2 suggest that the ACC plays a role in evaluating how much effort to expend for reward; however, the ACC is not necessary in all situations requiring an assessment of costs and benefits. In Experiment 3 we investigated learning and reversal learning of instrumental responses guided by reward predictive stimuli. A reaction time (RT) task demanding conditioned lever release was used in which the upcoming reward magnitude (five vs. one food pellet) was signalled in advance by discriminative visual stimuli. Results revealed that rats with ACC lesions were able to discriminate reward magnitude-predictive stimuli and to adapt instrumental behavior to reversed stimulus-reward magnitude contingencies. Thus, in a simple discrimination task as used here, the ACC appears not to be required to discriminate reward magnitude-predictive stimuli and to use the learned significance of the stimuli to guide instrumental behavior.     

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.     

Four‐month‐old infants individuate and track simple tools following functional demonstrations

Two experiments examined whether 4‐month‐olds (n = 120) who were induced to assign two objects to different categories would then be able to take advantage of these contrastive categorical encodings to individuate and track the objects. In each experiment, infants first watched functional demonstrations of two tools, a masher and tongs (Experiment 1) or a marker and a knife (Experiment 2). Next, half the infants saw the two tools brought out alternately from behind a screen, which was then lowered to reveal only one of the tools (different‐objects condition); the other infants saw similar events except that the same tool was shown on either side of the screen (same‐object condition). In both experiments, infants in the different‐objects condition looked reliably longer than those in the same‐object condition, and this effect was eliminated if the demonstrations involved similar but non‐functional actions. Together, these results indicate that infants (a) were led by the functional demonstrations they observed to assign the two tools to distinct categories, (b) recruited these categorical encodings to individuate and track the tools, and hence (c) detected a violation in the different‐objects condition when the screen was lowered to reveal only one tool. Categorical information thus plays a privileged role in individuation and identity tracking from a very young age.     

Making tools isn't child's play

Tool making evidences intelligent, flexible thinking. In Experiment 1, we confirmed that 4- to 7-year-olds chose a hook tool to retrieve a bucket from a tube. In Experiment 2, 3- to 5-year-olds consistently failed to innovate a simple hook tool. Eight-year-olds performed at mature levels. In contrast, making a tool following demonstration was easy for even the youngest children. In Experiment 3, children’s performance did not improve given the opportunity to manipulate the objects in a warm-up phase. Children’s tool innovation lags substantially behind their ability to learn how to make tools by observing others.     

Effects of reinforcer probability, delay, and response requirements on the choices of rats and pigeons: possible species differences

In Experiment 1 with rats, a left lever press led to a 5-s delay and then a possible reinforcer. A right lever press led to an adjusting delay and then a certain reinforcer. This delay was adjusted over trials to estimate an indifference point, or a delay at which the two alternatives were chosen about equally often. Indifference points increased as the probability of reinforcement for the left lever decreased. In some conditions with a 20% chance of food, a light above the left lever was lit during the 5-s delay on all trials, but in other conditions, the light was only lit on those trials that ended with food. Unlike previous results with pigeons, the presence or absence of the delay light on no-food trials had no effect on the rats' indifference points. In other conditions, the rats showed less preference for the 20% alternative when the time between trials was longer. In Experiment 2 with rats, fixed-interval schedules were used instead of simple delays, and the presence or absence of the fixed-interval requirement on no-food trials had no effect on the indifference points. In Experiment 3 with rats and Experiment 4 with pigeons, the animals chose between a fixed-ratio 8 schedule that led to food on 33% of the trials and an adjusting-ratio schedule with food on 100% of the trials. Surprisingly, the rats showed less preference for the 33% alternative in conditions in which the ratio requirement was omitted on no-food trials. For the pigeons, the presence or absence of the ratio requirement on no-food trials had little effect. The results suggest that there may be differences between rats and pigeons in how they respond in choice situations involving delayed and probabilistic reinforcers.     

Making choices in anticipation of similar future choices can increase self-control

Two experiments tested the efficacy of linking a current choice with similar future choices as a means of increasing self-control. Participants were offered choices between smaller and sooner vs. larger and later amounts of money (Experiment 1, n = 60) or food (Experiment 2, n = 34). After a small-large pair for which the participant preferred the smaller reward was found, a choice between the same pair was offered as the 1st of 5 such choices to be offered over a period of weeks. The majority of participants in both experiments who chose between all 5 smaller and all 5 larger rewards chose the larger rewards. One third of participants in Experiment 1 who could choose independently on each pair in the series reversed their previous preference and chose the larger reward in the context of the series. These results suggest that self-control can be enhanced by viewing one's current choice as predictive of future choices.     

Summation of symbols by pigeons (Columba livia): the importance of number and mass of reward items

In each of 3 experiments, different sets of 4 pigeons (Columba livia) were trained to discriminate between 2 visual symbols that covered wells containing food items that varied in number, mass, or both. In Experiment 1, the symbols were associated with 0, 1, 3, 5, 7, or 9 pieces of grain reward. The pigeons learned to choose the symbol corresponding to the larger reward, and on summation tests, they chose the pair of symbols that summed to the larger total reward. When number of food pellets was varied but mass of reward was held constant in Experiment 2, preference for the larger number symbols failed to appear. When number was held constant and mass was varied in Experiment 3, the pigeons showed a clear preference for the larger mass symbols on single-symbol and summation tests. These findings show that pigeons summate the value of symbols and are more likely to represent symbols by mass of food reward than by number of food items.     

Responses of observer rats (Rattus norvegicus) to complex, diet-related signals emitted by demonstrator rats

We explored the effects of complex, food-identifying signals emitted by demonstrator Long-Evans rats (Rattus norvegicus) on food preferences of their observers. In Experiments 1 and 2, observers identified each of 2 or 3 foods their demonstrators had eaten before interacting with observers. In Experiment 3, individual observers interacted with groups of demonstrators. Some of these demonstrators had eaten one food, some another. Observers then chose between the two foods. The greater the proportion of demonstrators in a group that had eaten a diet, the greater the proportion of that diet the observers ate. In Experiment 4, each observer interacted over several weeks with a series of demonstrators and preferred each of the foods its demonstrators had eaten. In sum, the food preferences of observers were affected by several different types of complex, food-identifying signals like those one might expect rats to encounter outside the laboratory.     

Relative sensitivity to reinforcer amount and delay in a self-control choice situation.

Rats were exposed to concurrent-chains schedules in which a single variable-interval schedule arranged entry into one of two terminal-link delay periods (fixed-interval schedules). The shorter delay ended with the delivery of a single food pellet; the longer day ended with a larger number of food pellets (two under some conditions and six under others). In Experiment 1, the terminal-link delays were selected so that under all conditions the ratio of delays would exactly equal the ratio of the number of pellets. But the absolute duration of the delays differed across conditions. In one condition, for example, rats chose between one pellet delayed 5 s and six pellets delayed 30 s; in another condition rats chose between one pellet delayed 10 s and six pellets delayed 60 s. The generalized matching law predicts indifference between the two alternatives, assuming that the sensitivity parameters for amount and delay of reinforcement are equal. The rats' choices were, in fact, close to indifference except when the choice was between one pellet delayed 5 s and six pellets delayed 30 s. That deviation from indifference suggests that the sensitivities to amount and delay differ from each other depending on the durations of the delays. In Experiment 2, rats chose between one pellet following a 5-s delay and six pellets following a delay that was systematically increased over sessions to find a point of indifference. Indifference was achieved when the delay to the six pellets was approximately 55 s. These results are consistent with the possibility that the relative sensitivities to amount and delay differ as a function of the delays.