Olfactory system receptor cells

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-... 

Eisthen H.L. (1992). Phylogeny of the vomeronasal system and of receptor cell-types in the olfactory and vomeronasal epithelia of vertebrates. Microsc Res Tech 23, 1-21. 

Even though well-characterized at a biophysical level, the mechanical transduction mechanism of hair cells is still not fully understood in molecular terms. This discrepancy is in part due to the extreme scarcity of hair cells instead of the millions or even hundreds of millions of receptor cells that the olfactory and visual systems possess, only a few tens of thousands of hair cells are found in the internal ears of most vertebrate species. The small number of hair cells and the direct transduction mechanism has greatly impeded molecular biological and... 

Differential sensory sensitivity. The insect s perception of plant odours differs essentially from their discrimination of non-volatile taste substances, as phytophagous insects may already perceive the odour at some distance from the plant. In adult phytophagous insects the antennae bear a large number of olfactory sensilla in order to detect the minute concentrations of the leaf odour components in the air downwind from a plant. The overall sensitivity of the antennal olfactory receptor system can be measured by making use of the electroantennogram technique (17). An electroantennogram (EAG) is the change in potential between the tip of an antenna and its base, in response to stimulation by an odour component. Such an EAG reflects the receptor potentials of the olfactory receptor cell population in the antenna. 

Recently, a putative olfactory receptor from Drosophila, Or43a (Clyne et al., 1999 Vosshall et al., 1999), has been expressed in Xenopus laevis oocytes (Wetzel et al., 2001). The receptor expressed in a heterologous cell system was activated by four odorants, i.e. cyclohexanone, cyclohexanol, benzaldehyde, and benzyl alcohol (Wetzel et al., 2001). These experiments not only provided direct evidence for the function of the Or gene, but also demonstrated that the olfactory receptor can be stimulated without an odorant-binding protein. It was demonstrated earlier that PBP was not necessary to obtain pheromone-dependent responses in cultured olfactory receptor neurons of Manduca sexta (Stengl et al., 1992). The possibility that OBPs have been produced in vitro and were present in cultured ORNs could not be excluded. The same argument can not be raised for the heterologous expression of the Drosophila olfactory receptor. While the evidence that Xenopus oocytes responded to odorants in the absence of OBPs does not support the OBP-odorant complex model, it also demonstrated that OBPs are essential for the kinetics of the olfactory system (see below). 

From the receptor cells nerves pass through the olfactory lobes at the front end of the brain direct to the central basal region, the part known as the "limbic system." This forms part of our deep-seated unconscious mind, being associated with the control of emotion and sexual activity, as well as with feelings of pleasure. In evolutionary terms it is also the oldest part of the brain, providing evidence of the early and continuing importance of the sense of smell in animal behavior. 

Figure 5.9 The human olfactory system. (A) Section through the nose. (B) Section through the cribriform plate. (C) The olfactory pathway to the cerebrum (forebrain). This shows the pathway of olfactory sensation. Nasal stimulation begins at the cilia of the olfactory receptor cells located at the ends of the olfactory nerves. The olfactory nerves then carry the impulse to the cerebrum, resulting in the sense of smell.

Of all these different steps, the very first one, namely the interaction of a stimulus, i.e. molecules that have a smell , with the actual receptor is not at all u nderstood. These receptors are supposed to be located in the membrane of the cilia cells, because these cilia are the furthest out of the antennaeof the olfactory system, and they have been shown to be excitable by chemical stimuli. 

Importantly, each taste receptor cell expresses many different members of the T2R family. This pattern of expression stands in sharp contrast to the pattern of one receptor type per cell that characterizes the olfactory system (Figure 32.16). The difference in expression patterns accounts for the much greater specificity of our perceptions of smells compared with tastes. We are able to distinguish among subtly different odors because each odorant stimulates a unique pattern of neurons. In contrast, many tastants stimulate the same neurons. Thus, we perceive only "bitter" without the ability to discriminate cycloheximide from quinine. 

---