The mysterious cognitive abilities of bees: why models of visual processing need to consider experience and individual differences in animal performance.

Vision is one of the most important modalities for the remote perception of biologically important stimuli. Insects like honeybees and bumblebees use their colour and spatial vision to solve tasks, such as navigation, or to recognise rewarding flowers during foraging. Bee vision is one of the most intensively studied animal visual systems, and several models have been developed to describe its function. These models have largely assumed that bee vision is determined by mechanistic hard-wired circuits, with little or no consideration for behavioural plasticity or cognitive factors. However, recent work on both bee colour vision and spatial vision suggests that cognitive factors are indeed a very significant factor in determining what a bee sees. Individual bumblebees trade-off speed for accuracy, and will decide on which criteria to prioritise depending upon contextual information. With continued visual experience, honeybees can learn to use non-elemental processing, including configural mechanisms and rule learning, and can access top-down information to enhance learning of sophisticated, novel visual tasks. Honeybees can learn delayed-matching-to-sample tasks and the rules governing this decision making, and even transfer learned rules between different sensory modalities. Finally, bees can learn complex categorisation tasks and display numerical processing abilities for numbers up to and including four. Taken together, this evidence suggests that bees do have a capacity for sophisticated visual behaviours that fit a definition for cognition, and thus simple elemental models of bee vision need to take account of how a variety of factors may influence the type of results one may gain from animal behaviour experiments.