Hooded crows (Corvus cornix) manufacture objects relative to a mental template.

It was recently found that not only tool-specialized New Caledonian crows, but also Goffin cockatoos can manufacture physical objects in accordance with a mental template. That is, they can emulate features of existing objects when they manufacture new items. Both species spontaneously ripped pieces of card into large strips if they had previously learned that a large template was rewarded, and small strips when they previously learned that a small template was rewarded. Among New Caledonian crows, this cognitive ability was suggested as a potential mechanism underlying the transmission of natural tool designs. Here, we tested for the same ability in another non-specialised tool user-Hooded crows (Corvus cornix). Crows were exposed to pre-made template objects, varying first in colour and then in size, and were rewarded only if they chose pre-made objects that matched the template. In subsequent tests, birds were given the opportunity to manufacture versions of these objects. All three crows ripped paper pieces from the same colour material as the rewarded template, and, crucially, also manufactured objects that were more similar in size to previously rewarded, than unrewarded, templates, despite the birds being rewarded at random in both tests. Therefore, we found the ability to manufacture physical objects relative to a mental template in yet another bird species not specialized in using or making foraging tools in the wild, but with a high level of brain and cognitive development.

A novel test of flexible planning in relation to executive function and language in young children.

In adult humans, decisions involving the choice and use of tools for future events typically require episodic foresight. Previous studies suggest some non-human species are capable of future planning; however, these experiments often cannot fully exclude alternative learning explanations. Here, we used a novel tool-use paradigm aiming to address these critiques to test flexible planning in 3- to 5-year-old children, in relation to executive function and language abilities. In the flexible planning task, children were not verbally cued during testing, single trials avoided consistent exposure to stimulus-reward relationships, and training trials provided experience of a predictable return of reward. Furthermore, unlike most standard developmental studies, we incorporated short delays before and after tool choice. The critical test choice included two tools with equal prior reward experience-each only functional in one apparatus. We tested executive function and language abilities using several standardized tasks. Our results echoed standard developmental research: 4- and 5-year-olds outperformed 3-year-olds on the flexible planning task, and 5-year-old children outperformed younger children in most executive function and language tasks. Flexible planning performance did not correlate with executive function and language performance. This paradigm could be used to investigate flexible planning in a tool-use context in non-human species.

Decision-making flexibility in New Caledonian crows, young children and adult humans in a multi-dimensional tool-use task.

The ability to make profitable decisions in natural foraging contexts may be influenced by an additional requirement of tool-use, due to increased levels of relational complexity and additional work-effort imposed by tool-use, compared with simply choosing between an immediate and delayed food item. We examined the flexibility for making the most profitable decisions in a multi-dimensional tool-use task, involving different apparatuses, tools and rewards of varying quality, in 3-5-year-old children, adult humans and tool-making New Caledonian crows (Corvus moneduloides). We also compared our results to previous studies on habitually tool-making orangutans (Pongo abelii) and non-tool-making Goffin's cockatoos (Cacatua goffiniana). Adult humans, cockatoos and crows, but not children and orangutans, did not select a tool when it was not necessary, which was the more profitable choice in this situation. Adult humans, orangutans and cockatoos, but not crows and children, were able to refrain from selecting non-functional tools. By contrast, the birds, but not the primates tested, struggled to attend to multiple variables-where two apparatuses, two tools and two reward qualities were presented simultaneously-without extended experience. These findings indicate: (1) in a similar manner to humans and orangutans, New Caledonian crows and Goffin's cockatoos can flexibly make profitable decisions in some decision-making tool-use tasks, though the birds may struggle when tasks become more complex; (2) children and orangutans may have a bias to use tools in situations where adults and other tool-making species do not.

Self-control in crows, parrots and nonhuman primates.

Self-control is critical for both humans and nonhuman animals because it underlies complex cognitive abilities, such as decision-making and future planning, enabling goal-directed behavior. For instance, it is positively associated with social competence and life success measures in humans. We present the first review of delay of gratification as a measure of self-control in nonhuman primates, corvids (crow family) and psittacines (parrot order): disparate groups that show comparable advanced cognitive abilities and similar socio-ecological factors. We compare delay of gratification performance and identify key issues and outstanding areas for future research, including finding the best measures and drivers of delayed gratification. Our review therefore contributes to our understanding of both delayed gratification as a measure of self-control and of complex cognition in animals. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Psychology > Comparative Psychology.

New Caledonian crows infer the weight of objects from observing their movements in a breeze.

Humans use a variety of cues to infer an object's weight, including how easily objects can be moved. For example, if we observe an object being blown down the street by the wind, we can infer that it is light. Here, we tested whether New Caledonian crows make this type of inference. After training that only one type of object (either light or heavy) was rewarded when dropped into a food dispenser, birds observed pairs of novel objects (one light and one heavy) suspended from strings in front of an electric fan. The fan was either on-creating a breeze which buffeted the light, but not the heavy, object-or off, leaving both objects stationary. In subsequent test trials, birds could drop one, or both, of the novel objects into the food dispenser. Despite having no opportunity to handle these objects prior to testing, birds touched the correct object (light or heavy) first in 73% of experimental trials, and were at chance in control trials. Our results suggest that birds used pre-existing knowledge about the behaviour exhibited by differently weighted objects in the wind to infer their weight, using this information to guide their choices.

Young children do not require perceptual-motor feedback to solve Aesop's Fable tasks.

Aesop's Fable tasks-in which subjects drop objects into a water-filled tube to raise the water level and obtain out-of-reach floating rewards -have been used to test for causal understanding of water displacement in both young children and non-human animals. However, a number of alternative explanations for success on these tasks have yet to be ruled out. One hypothesis is that subjects may respond to perceptual-motor feedback: repeating those actions that bring the reward incrementally closer. Here, we devised a novel, forced-choice version of the Aesop's Fable task to assess whether subjects can solve water displacement tasks when this type of feedback is removed. Subjects had to select only one set of objects, or one type of tube, into which all objects were dropped at once, and the effect the objects had on the water level was visually concealed. In the current experiment, fifty-five 5-9 year old children were tested in six different conditions in which we either varied object properties (floating vs. sinking, hollow vs. solid, large vs. small and too large vs. small objects), the water level (high vs. low) and/or the tube size (narrow vs. wide). We found that children aged 8-9 years old were able to solve most of the water displacement tasks on their first trial, without any opportunity for feedback, suggesting that they mentally simulated the results of their actions before making a choice. Children aged 5-7 years solved two conditions on their first trial (large vs. small objects and high- vs. low-water levels), and learnt to solve most of the remaining conditions over five trials. The developmental pattern shown here is comparable to previous studies using the standard Aesop's Fable task, where eight year olds are typically successful from their first trial and 5-7 year olds learn to pass over five trials. Thus, our results indicate that children do not depend on perceptual-motor feedback to solve these water displacement tasks. The forced-choice paradigm we describe could be used comparatively to test whether or not non-human animals require visual feedback to solve water displacement tasks.

Performance in Object-Choice Aesop's Fable Tasks Are Influenced by Object Biases in New Caledonian Crows but not in Human Children.

The ability to reason about causality underlies key aspects of human cognition, but the extent to which non-humans understand causality is still largely unknown. The Aesop's Fable paradigm, where objects are inserted into water-filled tubes to obtain out-of-reach rewards, has been used to test casual reasoning in birds and children. However, success on these tasks may be influenced by other factors, specifically, object preferences present prior to testing or arising during pre-test stone-dropping training. Here, we assessed this 'object-bias' hypothesis by giving New Caledonian crows and 5-10 year old children two object-choice Aesop's Fable experiments: sinking vs. floating objects, and solid vs. hollow objects. Before each test, we assessed subjects' object preferences and/or trained them to prefer the alternative object. Both crows and children showed pre-test object preferences, suggesting that birds in previous Aesop's Fable studies may also have had initial preferences for objects that proved to be functional on test. After training to prefer the non-functional object, crows, but not children, performed more poorly on these two object-choice Aesop's Fable tasks than subjects in previous studies. Crows dropped the non-functional objects into the tube on their first trials, indicating that, unlike many children, they do not appear to have an a priori understanding of water displacement. Alternatively, issues with inhibition could explain their performance. The crows did, however, learn to solve the tasks over time. We tested crows further to determine whether their eventual success was based on learning about the functional properties of the objects, or associating dropping the functional object with reward. Crows inserted significantly more rewarded, non-functional objects than non-rewarded, functional objects. These findings suggest that the ability of New Caledonian crows to produce performances rivaling those of young children on object-choice Aesop's Fable tasks is partly due to pre-existing object preferences.

Investigating animal cognition with the Aesop's Fable paradigm: Current understanding and future directions.

The Aesop's Fable paradigm - in which subjects drop stones into tubes of water to obtain floating out-of-reach rewards - has been used to assess causal understanding in rooks, crows, jays and human children. To date, the performance of corvids suggests that they can recognize the functional properties of a variety of objects including size, weight and solidity, and they seem to be more capable of learning from causal information than arbitrary information. However, 2 alternative explanations for their performance have yet to be ruled out. The perceptual-motor feedback hypothesis suggests that subjects may attend solely to the movement of the reward, repeating actions which bring the reward closer, while the object-bias hypothesis suggests that subjects could pass certain tasks by preferring to handle objects that resemble natural stones. Here we review our current understanding of performance on the Aesop's Fable tasks, and suggest that studies controlling for feedback and object preferences will help us determine exactly what animals understand about the cause and effect of water displacement.

New Caledonian crows rapidly solve a collaborative problem without cooperative cognition.

There is growing comparative evidence that the cognitive bases of cooperation are not unique to humans. However, the selective pressures that lead to the evolution of these mechanisms remain unclear. Here we show that while tool-making New Caledonian crows can produce collaborative behavior, they do not understand the causality of cooperation nor show sensitivity to inequity. Instead, the collaborative behavior produced appears to have been underpinned by the transfer of prior experience. These results suggest that a number of possible selective pressures, including tool manufacture and mobbing behaviours, have not led to the evolution of cooperative cognition in this species. They show that causal cognition can evolve in a domain specific manner-understanding the properties and flexible uses of physical tools does not necessarily enable animals to grasp that a conspecific can be used as a social tool.

Reasoning by exclusion in New Caledonian crows (Corvus moneduloides) cannot be explained by avoidance of empty containers.

Whether animals can reason or merely learn associatively is a long-standing debate. Researchers have approached this question by investigating whether dogs, birds, and primates can reason by exclusion (choosing by logically excluding all other alternatives). However, these studies have not resolved whether animals are capable of inferring which option is rewarded or are merely avoiding options known to be incorrect. Here, we used a forced-choice tubes task, where strategies of "reasoning by exclusion" and "avoidance of empty containers" predicted different responses. Two tubes (1 straight, 1 bent) were presented in 5 types of orientation, varying whether the rewarded location could be inferred. We compared predictions from both strategies with the observed performance of 8 wild-caught New Caledonian crows. Two of the 8 birds' choices were entirely consistent with reasoning by exclusion only. A further 4 birds followed a mixed strategy, where both reasoning and avoidance could have influenced their decisions. Thus, although avoidance plays a role, it cannot fully explain the crows' choices. Confirming how animals naturally solve problems is increasingly important in animal cognition; we demonstrate that NC crows can inferentially reason without explicit training, but, like humans, most do not rely solely on reasoning to make decisions.

Modifications to the Aesop's Fable paradigm change New Caledonian crow performances.

While humans are able to understand much about causality, it is unclear to what extent non-human animals can do the same. The Aesop's Fable paradigm requires an animal to drop stones into a water-filled tube to bring a floating food reward within reach. Rook, Eurasian jay, and New Caledonian crow performances are similar to those of children under seven years of age when solving this task. However, we know very little about the cognition underpinning these birds' performances. Here, we address several limitations of previous Aesop's Fable studies to gain insight into the causal cognition of New Caledonian crows. Our results provide the first evidence that any non-human animal can solve the U-tube task and can discriminate between water-filled tubes of different volumes. However, our results do not provide support for the hypothesis that these crows can infer the presence of a hidden causal mechanism. They also call into question previous object-discrimination performances. The methodologies outlined here should allow for more powerful comparisons between humans and other animal species and thus help us to determine which aspects of causal cognition are distinct to humans.

Using the Aesop's fable paradigm to investigate causal understanding of water displacement by New Caledonian crows.

Understanding causal regularities in the world is a key feature of human cognition. However, the extent to which non-human animals are capable of causal understanding is not well understood. Here, we used the Aesop's fable paradigm--in which subjects drop stones into water to raise the water level and obtain an out of reach reward--to assess New Caledonian crows' causal understanding of water displacement. We found that crows preferentially dropped stones into a water-filled tube instead of a sand-filled tube; they dropped sinking objects rather than floating objects; solid objects rather than hollow objects, and they dropped objects into a tube with a high water level rather than a low one. However, they failed two more challenging tasks which required them to attend to the width of the tube, and to counter-intuitive causal cues in a U-shaped apparatus. Our results indicate that New Caledonian crows possess a sophisticated, but incomplete, understanding of the causal properties of displacement, rivalling that of 5-7 year old children.