Molecular neurobiology of olfaction.

Odor discrimination is mediated via dendritic cilia of olfactory receptor neurons. Odorants traverse the aqueous mucous interphase that lines the surface of the olfactory neuroepithelium and interact with odorant receptors, which are members of the superfamily of G-protein-linked receptors. These interactions trigger synthesis of second messengers, including cyclic AMP and inositol triphosphate. Cyclic AMP opens a cation channel to elicit the generator current, which depolarizes the cell and, ultimately, leads to action potentials. Inositol triphosphate opens a calcium channel in the ciliary plasma membrane. Calcium entering through both this channel and the cyclic nucleotide-gated channel modulates the response to odorants by amplifying the generation of cyclic AMP after binding to calmodulin. Calcium also is essential for desensitization of olfactory receptor neurons. Differential expression of odorant receptors of diverse ligand specificities by different olfactory neurons ensures that the structures and concentrations of odorants that reach the chemosensory surface are encoded as distinct patterns of neuronal activity, which are relayed to the brain where they take shape as characteristic odor sensations.