Olfactory system epithelium

Odors are perceived via the olfactory system, which is composed of two organs in the nose the olfactory epithelium, a very small area in the nasal system, and the trigeminal nerve endings, which are much more widely distributed in the nasal cavity (11). The olfactory epithelium is extremely sensitive, and humans often sniff to bring more odorant in contact with this area. The trigeminal nerves initiate protective reflexes, such as sneezing or interruption of irrhalation, with exposure to noxious odorants. 

The mechanisms by which the taste (and also the olfactory) system senses chemical compounds is assumed to occur by way of a chemoreceptory system that interacts effectively with a broad, structural variety of stimulant molecules, by means of a receptor epithelium consisting of the mosaic of adjacent, peripheral membranes of many receptor cells, exposed to a medium carrying stimulus molecules. A receptor cell is conveniently and, for our present purpose, sufficiently defined as a cell equipped to interact, according to some mechanism, with stimulus molecules, to convert the effect of this interaction into a signal, and to project this signal into the system. The taste receptor is thus a differentiated, epithelial cell synaptically contact-... 

Two olfactory systems have evolved in terrestrial vertebrates which differ in both their peripheral anatomy and central projections. The main olfactory system is usually conceived as a general analyzer that detects and differentiates among complex chemosignals of the environment (Firestein 2001). Odors are detected by olfactory sensory neurons located in the main olfactory epithelium (MOE) these neurons project to glomeruli in the main olfactory bulb (MOB). The mitral and tufted neurons abutting these MOB glomeruli then transmit olfactory signals to various... 

As with many macrosmatic mammals, rodents have two separate chemosensory systems, the main olfactory system (MOS) and accessory olfactory system (AOS), which respond to social odors. Importantly, these sensory systems differ not only in their peripheral morphology and central projections, but also in the types of chemosignals that they process (Meredith 1991). Sensory neurons of the MOS, which are located in the main olfactory epithelium and project to the main olfactory bulbs, process volatile chemicals and can detect odors at a distance. In contrast, sensory neurons of the AOS, which are located in the vomeronasal organs (VNO) and project to the accessory olfactory bulbs, primarily process large, non-volatile chemicals and require contact for stimulation (Meredith 1991). 

Pioneering efforts to understand the nature of olfactory coding were reported by Adrian (24-27). His work introduced the ideas that different odors activate ORCs in different regions of the olfactory epithelium and that spatiotemporal patterns of ORC firing would suffice to encode different odors. Subsequent studies by many investigators and involving various recording methods (reviewed in refs. 13 and 28) led to the conclusion that, at various levels of the pathway, the olfactory system uses distributed neural activity to encode information about olfactory stimuli. 

It is not always clear how pheromone signals are detected in mammals. Most vertebrates, mice for example, have a VNO in addition to the main olfactory system. The VNO has two separate families of olfactory receptors Vlr, 137 functional receptors in mice V2r, 60 functional receptors in mice. The genes for these are only distantly related to those for the main olfactory receptors, suggesting that these systems evolved independently. As a general rule, it is the VNO and not the olfactory epithelium that is responsible for detecting pheromone molecules. However, it has been demonstrated that mice whose VNO has been surgically removed can discriminate MHC-determined odor types. This finding clearly implicates the main olfactory system in the detection of pheromones. 

In mammals, the main olfactory system is the work horse in the perception of odors. Excellent detailed reviews of the mammalian olfactory system are available elsewhere. In brief, the olfactory epithelium is located on a portion of the scroll bones (endoturbinales and posterior part of nasoturbinales F ig. 5.4), in humans it is located about 1 cm beneath the bridge of the nose. Olfactory reception is affected by several factors, such as the size, shape, or wetness of the nasal passages. In the dog, the olfactory membrane extends over 75-150 cm depending on body size, while in humans it is only 2-4cm. It consists of three... 

Although the vomeronasal system is specialized to detect stimuli in a liquid environment, it probably is not functional in utero, at least in mice. Fluorescent microspheres were not taken up by the vomeronasal organ as the access canal is not open yet in utero. In rats, by contrast, the canal is open before birth and the microspheres can be taken up. The olfactory epithelium of the main olfactory system plays a greater role prenatally, as evidenced by the uptake of radiolabeled 2-deoxyglucose (Coppola and Coltrane 1994). Fetal mice respond to amyl acetate and isovaleric acid delivered into the nasal cavity through a tiny cannula (Coppola, 2001). In both rats and mice, the main olfactory system, and not the vomeronasal system, appears to mediate prenatal olfaction (Coppola, 2001). 

Shipley MT Transport of molecules from nose to brain transneuronal anterograde and retrograde labeling in the rat olfactory system by wheat germ agglutinin-horseradish peroxidase applied to the nasal epithelium. Brain Res Bull 15 129-142, 1985... 

Is the peripheral olfactory system of insects composed of essentially separate units or can it be considered a continuous compartmentalized epithelium such as the insect eye Evidence from studies on developmental mechanisms suggests it is a bit of both. 

Olfaction, like visual and taste perception, is an ancient process. Olfaction plays a role in sexual arousal. The olfactory system in mammals is remarkable with respect to the number of receptors engaged in monitoring odours. There are several thousand hepta-helical G-protein-coupled receptors in the olfactory epithelium and the nasal organ of a dog, and still about 1000 receptors in the corresponding human organs. It has been estimated that nearly 1% of all genes code for olfactory receptors alone. 

Fig. 2 Axonal projection in the mouse olfactory system, a The transgenic mouse, MOR28-IRES-tau-lacZ (Serizawa et al. 2000), stained with X-gal (lateral view). Olfactory sensory neurons (OSNs) expressing the taw-/acZ-tagged MOR28 gene project their axons to a specific site forming a glomerulus in the olfactory bulb (OB), b Spatial correlation between the olfactory epithelium (OE) and the OB in the mouse olfactory system. OSNs in the dorsomedial zone (D zone) in the OE project their axons to the dorsal domain (D domain) of the OB. Class I ORs are mostly expressed by OSNs in the D zone in the OE, which target the anterodorsal cluster of the D domain in the OB. In the ventrolateral zone (V zone), each class II OR possesses its own unique expression...

Ressler KJ, Sullivan SL, Buck LB (1993) A zonal organization of odorant receptor gene expression in the olfactory epithelium.Cell 73(3) 597-609 Ressler KJ, Sullivan SL, Buck LB (1994) Information coding in the olfactory system evidence for a stereotyped and highly organized epitope map in the olfactory bulb.Cell 79(7) 12451255... 

Fig. 20. Olfactory epithelium projections to the MOB. Photomicrographs of sagittal sections through the olfactory bulb In sections stained for Nissl (A) or with WGA HRP after injection of the tracer in the olfactory epithelium (B). Note that most of the olfactory bulb is comprised by the main olfactory system while a small portion of the dorsocaudal bulb is occupied by the accessory olfactory bulb in the rat. Note also in B that the WGA HRP did not transport to the glomeruli of AOB since the tracer did not gain access to the vomeronasal organ that is embedded in the nasal septum. Bar in B, 1 mm.

The peripheral organ for the olfactory system is the olfactory sensory epithelium, located in the dorsal-posterior portion of the nasal cavity. This epithelium, composed of several different types of cells, contains bipolar neurons (also known as olfactory receptor cells) whose axons extend to the brain and terminate in the main olfactory bulb. The VN organ is also situated in the periphery, and similarly contains several different types of cells, the most numerous being bipolar neurons whose axons terminate in the accessory olfactory bulb. The VN organ opens into the roof of the mouth via a very narrow channel, the VN duct. In snakes the tongue delivers odorants to the opening of the VN duct and, by a mechanism at present not understood, these odorants reach the dendritic tips of the bipolar neurons (Halpern and Kubie, 1980). 

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