Non-elemental processing in olfactory discrimination tasks needs bilateral input in honeybees

In patterning discriminations, animals have to differentiate a compound stimulus AB from each of its elements A and B. In positive patterning (PP), the compound is reinforced whilst the single elements are non-reinforced. In negative patterning (NP), single elements are reinforced whilst the compound is non-reinforced. Using olfactory conditioning of the proboscis extension response (PER), we asked whether honeybees (Apis mellifera) can solve these patterning problems when odorants are given unilaterally as well as bilaterally to the antennae. Separating the olfactory input space of bees into two independent zones using plastic walls placed between the antennae, we conditioned bees in PP and NP procedures, with input on one side, on both sides, or in an ambiguous problem where bees had to solve PP on one side and NP on the other side. We found that bees with simultaneous bilateral input solve both patterning tasks efficiently. In contrast, PP but not NP was learned by bees receiving unilateral olfactory input. Bees subjected to the ambiguous NP/PP problem only solved PP. As PP can be solved through mere elemental processes, but NP is critically dependent on the use of non-elemental learning processes, our results suggest that bilateral olfactory input is necessary for non-elemental processing to take place in the bee brain.