Olfaction

Olfaction is a chemoreception that forms the sense of smell. Olfaction has many purposes, such as the detection of hazards, pheromones, and food. It integrates with other senses to form the sense of flavor. Olfaction is a chemoreception that forms the sense of smell. Olfaction has many purposes, such as the detection of hazards, pheromones, and food. It integrates with other senses to form the sense of flavor. Olfaction occurs when odorants bind to specific sites on olfactory receptors located in the nasal cavity. Glomeruli aggregate signals from these receptors and transmit them to the olfactory bulb, where the sensory input will start to interact with parts of the brain responsible for smell identification, memory, and emotion. Often, land organisms will have separate olfaction systems for smell and taste (orthonasal smell and retronasal smell), but water-dwelling organisms usually have only one system. In vertebrates, smells are sensed by olfactory sensory neurons in the olfactory epithelium. The olfactory epithelium is made up of at least six morphologically and biochemically different cell types. The proportion of olfactory epithelium compared to respiratory epithelium (not innervated, or supplied with nerves) gives an indication of the animal's olfactory sensitivity. Humans have about 10 cm2 (1.6 sq in) of olfactory epithelium, whereas some dogs have 170 cm2 (26 sq in). A dog's olfactory epithelium is also considerably more densely innervated, with a hundred times more receptors per square centimeter. Molecules of odorants passing through the superior nasal concha of the nasal passages dissolve in the mucus that lines the superior portion of the cavity and are detected by olfactory receptors on the dendrites of the olfactory sensory neurons. This may occur by diffusion or by the binding of the odorant to odorant-binding proteins. The mucus overlying the epithelium contains mucopolysaccharides, salts, enzymes, and antibodies (these are highly important, as the olfactory neurons provide a direct passage for infection to pass to the brain). This mucus acts as a solvent for odor molecules, flows constantly, and is replaced approximately every ten minutes. In insects, smells are sensed by olfactory sensory neurons in the chemosensory sensilla, which are present in insect antenna, palps, and tarsa, but also on other parts of the insect body. Odorants penetrate into the cuticle pores of chemosensory sensilla and get in contact with insect odorant-binding proteins (OBPs) or Chemosensory proteins (CSPs), before activating the sensory neurons. The binding of the ligand (odor molecule or odorant) to the receptor leads to an action potential in the receptor neuron, via a second messenger pathway, depending on the organism. In mammals, the odorants stimulate adenylate cyclase to synthesize cAMP via a G protein called Golf. cAMP, which is the second messenger here, opens a cyclic nucleotide-gated ion channel (CNG), producing an influx of cations (largely Ca2+ with some Na+) into the cell, slightly depolarising it. The Ca2+ in turn opens a Ca2+-activated chloride channel, leading to efflux of Cl−, further depolarizing the cell and triggering an action potential. Ca2+ is then extruded through a sodium-calcium exchanger. A calcium-calmodulin complex also acts to inhibit the binding of cAMP to the cAMP-dependent channel, thus contributing to olfactory adaptation. The main olfactory system of some mammals also contains small subpopulations of olfactory sensory neurons that detect and transduce odors somewhat differently. Olfactory sensory neurons that use trace amine-associated receptors (TAARs) to detect odors use the same second messenger signaling cascade as do the canonical olfactory sensory neurons. Other subpopulations, such as those that express the receptor guanylyl cyclase GC-D (Gucy2d) or the soluble guanylyl cyclase Gucy1b2, use a cGMP cascade to transduce their odorant ligands. These distinct subpopulations (olfactory subsystems) appear specialized for the detection of small groups of chemical stimuli. This mechanism of transduction is somewhat unusual, in that cAMP works by directly binding to the ion channel rather than through activation of protein kinase A. It is similar to the transduction mechanism for photoreceptors, in which the second messenger cGMP works by directly binding to ion channels, suggesting that maybe one of these receptors was evolutionarily adapted into the other. There are also considerable similarities in the immediate processing of stimuli by lateral inhibition.