Olfactory receptor neurons transduction

Fadool, D. A. and Ache, B. W., Plasma membrane inositol 1,4,5-trisphosphate-activated channels mediate signal transduction in lobster olfactory receptor neurons, Neuron, 9, 907, 1992. 

As is the case for all sensory pathways, the capacity to perceive and respond to olfactory cues (odorants) is the combined result of events that take place in both peripheral and central processing centers. These steps, which will be discussed in detail below, begin with the molecular transduction of chemical signals in the form of odorants into electrical activity by olfactory receptor neurons (ORNs) in the periphery whose axonal projections form characteristic synaptic connections with elements of the central nervous system (CNS). Within the CNS, complex patterns of olfactory signals are integrated and otherwise processed to afford recognition and ultimately, the behavioral responses to the insect s chemical environment. Within the context of pheromone recognition these responses would likely be centered on various elements of the insect s reproductive cycle. 

What happens at the instant when an odor-active molecule comes in touch with our nasal cavity 205 The first interaction of odorant molecules takes place in the olfactory receptor neurons, which are embedded in the pseudostratified columnar epithelium (or simply, olfactory epithelium), which is located in the posterior nasal cavity in the case of mammals. Olfactory sensory neurons express receptor proteins on the surface membrane of the cilia, which gain access to the extracellular region covered with mucus. The airborne odorants are dissolved into the mucus, bind with the receptors, and then the receptor protein triggers a signal transduction cascade. This results in the opening of the cation channel that would depolarize the sensory neuron and eventually elicit a train of action potentials in the axon. The olfactory axon leads to the olfactory bulb through basal lamina and lamina propria. 

Hansen A, Rolen SH, Anderson K, Morita Y, Caprio J, Finger TE (2003) Correlation between olfactory receptor cell type and function in the channel catfish. J Neurosci 23 347-359 Hansen A, Anderson KT, Finger TE (2004) Differential distribution of olfactory receptor neurons in goldfish structural and molecular correlates. J Comp Neurol 477 347-359 Hansen A, Zielinski BS (2005) Diversity in the olfactory epithelium of bony fishes development, lamellar arrangement, sensory neuron cell types and transduction components. J Neurocytol 34 183-208... 

Michel WC (1999) Cyclic nucleotide-gated channel activation is not required for activity-dependent labeling of zebrafish olfactory receptor neurons by amino adds. Biol Signals Recept 8 338-347 Michel WC, Derbidge DS (1997) Evidence of distinct amino acid and bile salt receptors in the olfactory system of the zebrafish, Danio rerio. Brain Res 764 179-187 Michel WC, Sanderson MJ, Olson JK, Lipschitz DL (2003) Evidence of a novel transduction pathway mediating detection of polyamines by the zebrafish olfactory system. J Exp Biiol 206 1697-1706... 

Fig. 4 The mechanism of insect olfactory transduction. Odorants and pheromones regulate the channel opening probability of the insect OR-Or83b receptor complex. An influx of cations depolarizes the olfactory receptor neuron (ORN), resulting in an increase in the firing rate. Depression of channel activity suppresses neural electrical activity. See Sato et al. (2008) for more details...

Breer H., Sense of smell Signal recognition and transduction in olfactory receptor neurons, in Handbook of Biosensors and Electronic Noses Medicine, Food and Environment, ed. E. Kress-Rogers (Boca Raton, EL CRC Press, 1997, 521-532). 

Mammalian pheromones released into their environment can readily reach their target tissue, either the main olfactory epithelium (MOE) or the VNO. Both target tissues are lined with an olfactory neuroepithelium that contains membrane-bound receptor proteins, which comprise the largest known family of G-protein-coupled [262] receptors in mammals. The number of mammalian olfactory receptors [263-265] found has been astonishing, but not unreasonable. The MOE and VNO have some common features, but also significant difierences in neuron types, primary structures of receptor proteins and signal transduction [266]. 

The 10 20 million olfactory sensory neurons in the human nose are confined to a relatively small patch of tissue located high in the nasal cavity. When odorants, e.g., carvone in Fig. 2, are deposited within the mucus covering the distal ends of the olfactory receptor cells, they interact with some of the membrane-bound, G-protein-coupled receptors [9]. This interaction initiates a transduction process that converts physicochemical information in the odorant, e.g., its structure or other attributes, into electrical energy that is conveyed in the form of pulses (action potentials) along olfactory axons to the brain. 

Figure 2 Depiction of some components of the vertebrate olfactory epithelium in the nose. Odorants, e.g., carvone, deposit themselves in the mucous layer and interact with molecular receptors in the membrane of cilia of the olfactory receptor cells. Subsequent to intracellular signal transduction events, action potentials are sent via the olfactory axons to the olfactory bulbs in the brain. Supporting cells provide physical and physiological support for the olfactory neurons. Undifferentiated basal (stem) cells are the source of new supporting and olfactory receptor cells.

The taste cells are situated in the lingual epithelium with the apical membrane exposed to the mucosal surface of the oral cavity and the basal surface in contact with the nerve [interstitial fluid] [FIGURE 10]. Within the basolateral surface are the nerves which respond to the chemestiietic stimulants, i.e. direct nerve stimulation. The microvilli at the apical membrane contain receptor proteins which respond to sweeteners, some bitters and possibly coolants. The olfactory cells are bipolar neurons with dendritic ends containing cilia exposed to the surface and axons linked to the brain, where they synapse in the olfactory bulb. The transfer of information from this initial stimulus-receptor interaction to the brain processing centers involves chentical transduction steps in the membrane and within the receptor cells. The potential chemical interactions at the cell membrane and within the cell are schematically outlined in FIGURE 10. 

Some olfactory neurons may use a second transduction mechanism. They have receptors coupled through G proteins to PLC rather than to adenylyl cyclase. Signal reception in these cells triggers production of IP3 (Fig. 12-19), which opens IP3-gated Ca2+ channels in the ciliary membrane. Influx of Ca2+ then depolarizes the ciliary membrane and generates a receptor potential or regulates Ca2+-dependent enzymes in the olfactory pathway. 

Ache and coworkers demonstrated that both cyclic nucleotides and inositol phosphates mediate the transduction of environmental chemical signals by the olfactory neurons of P. argus.62 65 Both biochemical and molecular biological techniques have shown that the receptor cells contain various G-protein subunits that would be necessary for signal detection by G-protein-associated chemoreceptors.48 49 66-69 In combination with electrophysiological studies,... 

Very recent efforts to further evaluate the functional implications of phosphoinositol signaling in vertebrate chemosensory transduction have led to the discovery that not only the phosphoinositol breakdown pathway catalyzed by phospholipase C, but also membrane phosphoinositols themselves involving phosphoinositide-3-kinase may play a functional role in the transduction process. It was found that 3-phosphoinositides signaling in concert with the canonical phosphoinositide turnover pathway modulate the cyclic nucleotide signaling cascade downstream of the receptor. The data suggest that 3-phosphoinositide, the primary product of PI3K activity, attenuates the cyclic nucleotide-dependent excitation of olfactory neurons (Spehr et al., 2002). 

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