Bird brains and tool use: beyond instrumental conditioning.

Few displays of complex cognition are as intriguing as nonhuman tool use. Long thought to be unique to humans, evidence for tool use and manufacture has now been gathered in chimpanzees, dolphins, and elephants. Outside of mammals, tool use is most common in birds, especially in corvids and parrots. The present paper reviews the evidence for avian tool use, both in the wild and in laboratory settings. It also places this behavioral evidence in the context of longstanding debates about the kinds of mental processes nonhumans can perform. Descartes argued that animals are unable to think because they are soulless machines, incapable of flexible behavior. Later, as human machines became more sophisticated and psychologists discovered classical and instrumental conditioning, skepticism about animal thinking decreased. However, behaviors that involve more than simple conditioning continued to elicit skepticism, especially among behaviorists. Nonetheless, as reviewed here, strong behavioral data now indicate that tool use in some birds cannot be explained as resulting entirely from instrumental conditioning. The neural substrates of tool use in birds remain unclear, but the available data point mainly to the caudolateral nidopallium, which shares both functional and structural features with the mammalian prefrontal cortex. As more data on the neural mechanisms of complex cognition in birds accrue, skepticism about those mental capacities should continue to wane.