In Search for the Retrievable Memory Trace in an Insect Brain

The search strategy for the memory trace and its semantics is exemplified for the case of olfactory learning in the honeybee brain. The logic of associative learning is used to guide the experimental approach into the brain by identifying the anatomical and functional convergence sites of the conditioned stimulus and unconditioned stimulus pathways. Two of the several convergence sites are examined in detail, the antennal lobe as the first-order sensory coding area, and the input region of the mushroom body as a higher order integration center. The memory trace is identified as the pattern of associative changes on the level of synapses. The synapses are recruited, drop out, and change the transmission properties for both specifically associated stimulus and the non-associated stimulus. Several rules extracted from behavioral studies are found to be mirrored in the patterns of synaptic change. The strengths and the weaknesses of the honeybee as a model for the search for the memory trace are addressed in a comparison with Drosophila. The question is discussed whether the memory trace exists as a hidden pattern of change if it is not retrieved and whether an external reading of the content of the memory trace may ever be possible. Doubts are raised on the basis that the retrieval circuits are part of the memory trace. The concept of a memory trace existing beyond retrieval is defended by referring to two well-documented processes also in the honeybee, memory consolidation during sleep, and transfer of memory across brain areas.