A ubiquitous feature of an animal's response to an odorant is that it declines when the odorant is frequently or continuously encountered. This decline in olfactory response, termed olfactory habituation, can have temporally or mechanistically different forms. The neural circuitry of the fruit fly Drosophila melanogaster's olfactory system is well defined in terms of component cells, which are readily accessible to functional studies and genetic manipulation. This makes it a particularly useful preparation for the investigation of olfactory habituation. In addition, the insect olfactory system shares many architectural and functional similarities with mammalian olfactory systems, suggesting that olfactory mechanisms in insects may be broadly relevant. In this chapter, we discuss the likely mechanisms of olfactory habituation in context of the participating cell types, their connectivity, and their roles in sensory processing. We overview the structure and function of key cell types, the mechanisms that stimulate them, and how they transduce and process odor signals. We then consider how each stage of olfactory processing could potentially contribute to behavioral habituation. After this, we overview a variety of recent mechanistic studies that point to an important role for potentiation of inhibitory synapses in the primary olfactory processing center, the antennal lobe, in driving the reduced response to familiar odorants. Following the discussion of mechanisms for short- and long-term olfactory habituation, we end by considering how these mechanisms may be regulated by neuromodulators, which likely play key roles in the induction, gating, or suppression of habituated behavior, and speculate on the relevance of these processes for other forms of learning and memory.
Keywords: Drosophila; antennal lobe; habituation; learning; memory; neuromodulator; olfactory habituation; recurrent inhibition; synaptic plasticity; translational control.
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