A cognitive account of trace conditioning in insects

Trace conditioning is a form of Pavlovian learning in which the conditioned stimulus (CS) and the unconditioned stimulus (US) are separated by a temporal gap. Insects learn trace associations of variable nature (appetitive, aversive) and involving CSs of different sensory modalities (olfactory, visual). The accessibility of the insect neural system in behaving animals allowed identifying neural processes driving trace conditioning: the existence of prolonged neural responses to the CS after stimulus offset and the anticipation of US responses during the free-stimulus gap. Specific brain structures, such as the mushroom bodies seem to be allocated to this learning form. Here, we posit that a further component facilitating trace conditioning in insects relates to neuromodulatory mechanisms underlying enhanced attention. We thus propose a model based on different types of mushroom-body neurons, which provides a cognitive account of trace conditioning in insects.