Cognitive neuroethology: dissecting non-elemental learning in a honeybee brain.

The brain of a honeybee contains only 960,000 neurons and its volume represents only 1 mm3. However, it supports impressive behavioral capabilities. Honeybees are equipped with sophisticated sensory systems and have well developed learning and memory capacities, whose essential mechanisms do not differ drastically from those of vertebrates. Here, I focus on non-elemental forms of learning by honeybees. I show that bees exhibit learning abilities that have been traditionally ascribed to a restricted portion of vertebrates, as they go beyond simple stimulus-stimulus or response-stimulus associations. To relate these abilities to neural structures and functioning in the bee brain we focus on the antennal lobes and the mushroom bodies. We conclude that there is a fair chance to understand complex behavior in bees, and to identify the potential neural substrates underlying such behavior by adopting a cognitive neuroethological approach. In such an approach, behavioral and neurobiological studies are combined to understand the rules and mechanisms of plastic behavior in a natural context.