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Humans are capable of simply observing a correlation between cause and effect, and then producing a novel behavioural pattern in order to recreate the same outcome. However, it is unclear how the ability to create such causal interventions evolved. Here, we show that while 24-month-old children can produce an effective, novel action after observing a correlation, tool-making New Caledonian crows cannot. These results suggest that complex tool behaviours are not sufficient for the evolution of this ability, and that causal interventions can be cognitively and evolutionarily disassociated from other types of causal understanding., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)

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.

Animals rarely solve problems spontaneously. Some bird species, however, can immediately find a solution to the string-pulling problem. They are able to rapidly gain access to food hung on the end of a long string by repeatedly pulling and then stepping on the string. It is currently unclear whether these spontaneous solutions are produced by insight or by a perceptual-motor feedback loop. Here, we presented New Caledonian crows and humans with a novel horizontal string-pulling task. While the humans succeeded, no individual crow showed a significant preference for the connected string, and all but one failed to gain the food even once. These results clearly show that string pulling in New Caledonian crows is generated not by insight, but by perceptual feedback. Animals can spontaneously solve problems without planning their actions.

The ability to make inferences about hidden causal mechanisms underpins scientific and religious thought. It also facilitates the understanding of social interactions and the production of sophisticated tool-using behaviors. However, although animals can reason about the outcomes of accidental interventions, only humans have been shown to make inferences about hidden causal mechanisms. Here, we show that tool-making New Caledonian crows react differently to an observable event when it is caused by a hidden causal agent. Eight crows watched two series of events in which a stick moved. In the first set of events, the crows observed a human enter a hide, a stick move, and the human then leave the hide. In the second, the stick moved without a human entering or exiting the hide. The crows inspected the hide and abandoned probing with a tool for food more often after the second, unexplained series of events. This difference shows that the crows can reason about a hidden causal agent. Comparative studies with the methodology outlined here could aid in elucidating the selective pressures that led to the evolution of this cognitive ability.

New Caledonian crows were presented with Bird and Emery's (2009a) Aesop's fable paradigm, which requires stones to be dropped into a water-filled tube to bring floating food within reach. The crows did not spontaneously use stones as tools, but quickly learned to do so, and to choose objects and materials with functional properties. Some crows discarded both inefficient and non-functional objects before observing their effects on the water level. Interestingly, the crows did not learn to discriminate between functional and non-functional objects and materials when there was an arbitrary, rather than causal, link between object and reward. This finding suggests that the crows' performances were not based on associative learning alone. That is, learning was not guided solely by the covariation rate between stimuli and outcomes or the conditioned reinforcement properties acquired by functional objects. Our results, therefore, show that New Caledonian crows can process causal information not only when it is linked to sticks and stick-like tools but also when it concerns the functional properties of novel types of tool.

The ability of some bird species to pull up meat hung on a string is a famous example of spontaneous animal problem solving. The "insight" hypothesis claims that this complex behaviour is based on cognitive abilities such as mental scenario building and imagination. An operant conditioning account, in contrast, would claim that this spontaneity is due to each action in string pulling being reinforced by the meat moving closer and remaining closer to the bird on the perch. We presented experienced and naïve New Caledonian crows with a novel, visually restricted string-pulling problem that reduced the quality of visual feedback during string pulling. Experienced crows solved this problem with reduced efficiency and increased errors compared to their performance in standard string pulling. Naïve crows either failed or solved the problem by trial and error learning. However, when visual feedback was available via a mirror mounted next to the apparatus, two naïve crows were able to perform at the same level as the experienced group. Our results raise the possibility that spontaneous string pulling in New Caledonian crows may not be based on insight but on operant conditioning mediated by a perceptual-motor feedback cycle.