Learning is an animal’s capacity to change behaviour, as the result of individual experience, so that it is better adapted to the animal’s physical and social environments. Learning helps to fine-tune behaviour in flexible ways that would be difficult to encode genetically. It is an inherent property of nervous systems in even the simplest animals. Memory is an animal’s capacity to retain learned information to influence future behaviour. ‘Learning and animal culture’ discusses both classical and operant conditioning; innate responses that do not require learning; social learning, which across generations leads to cultural traditions; and the use of tools, which is now known not to be a uniquely human ability.