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INTRODUCTION Smell and taste are generally classified as visceral senses because of their close association with gastrointestinal function. Physiologically, they are related to each other. The flavors of various foods are in large part a combination of their taste and smell. Consequently, food may taste "different" if one has a cold that depresses the sense of smell. Both taste and smell receptors are chemoreceptors that are stimulated by molecules in solution in mucus in the nose and saliva in the mouth. However, these two senses are anatomically quite different. The smell receptors are distance receptors (teleceptors); the smell pathways have no relay in the thalamus; and there is no neocortical projection area for olfaction. The taste pathways pass up the brain stem to the thalamus and project to the postcentral gyrus along with those for touch and pressure sensibility from the mouth. SMELL Olfactory Mucous Membrane The olfactory receptor cells are located in a specialized portion of the nasal mucosa, the yellowish-pigmented olfactory mucous membrane. In humans, it covers an area of 5 cm2 in the roof of the nasal cavity near the septum. It contains supporting cells and progenitor cells for the olfactory receptors. Interspersed between these cells are 10-20 million receptor cells. Each olfactory receptor is a neuron, and the olfactory mucous membrane is said to be the place in the body where the nervous system is closest to the external world. Each neuron has a short, thick dendrite with an expanded end called an olfactory rod. From these rods, cilia project to the surface of the mucus. The cilia are unmyelinated processes about 2 um long and 0.1 um in diameter. There are 10-20 cilia per receptor neuron. The axons of the olfactory receptor neurons pierce the cribriform plate of the ethmoid bone and enter the olfactory bulbs. The olfactory neurons, like the taste receptor cells but unlike most other neurons, are constantly being replaced with a half-time of a few weeks. The olfactory renewal process is carefully regulated, and there is evidence that in this situation, a bone morphogenic protein (BMP) exerts an inhibitory effect. The olfactory mucous membrane is constantly covered by mucus. This mucus is produced by Bowman's glands, which are just under the basal lamina of the membrane. Olfactory Bulbs In the olfactory bulbs, the axons of the receptors contact the primary dendrites of the mitral cells and tufted cells to form the complex globular synapses called olfactory glomeruli. The tufted cells are smaller than the mitral cells and have thinner axons, but both types send axons into the auditory cortex and the rest of the brain, and they appear to be similar from a functional point of view. An average of 26,000 receptor cell axons converges on each glomerulus. In addition to mitral and tufted cells, the olfactory bulbs contain periglomerular cells, which are inhibitory neurons connecting one glomerulus to another, and granule cells, which have no axons and make reciprocal synapses with the lateral dendrites of the mitral and tufted cells.
At these synapses, the mitral or tufted cell excites the granule cell by releasing glutamate,
and the granule cell side of the synapse in turn inhibits the mitral or tufted cell by releasing GABA. Olfactory Cortex The axons of the mitral and tufted cells pass posteriorly through the intermediate olfactory stria and the lateral olfactory stria to the olfactory cortex. The axons terminate on the apical dendrites of pyramidal cells in the olfactory cortex. In humans, sniffing activates the piriform cortex, but smells with or without sniffing activates the lateral and anterior orbitofrontal gyri of the frontal lobe. The orbitofrontal activation is generally greater on the right side than the left. Thus, the cortical representation of olfaction is asymmetric. Other fibers project to the amygdala, which is probably involved with the emotional responses to olfactory stimuli, and to the entorhinal cortex, which is concerned with olfactory memories.
Olfactory Thresholds & Discrimination Olfactory receptors respond only to substances that are in contact with the olfactory epithelium and are dissolved in the thin layer of mucus that covers it. The olfactory thresholds for the representative substances illustrate the remarkable sensitivity of the olfactory receptors to some substances. Methyl mercaptan, one of the substances in garlic, can be smelled at a concentration of less than 500 pg/L of air. In addition, olfactory discrimination is remarkable; for example, humans can recognize more than 10,000 different odors. Determination of differences in the intensity of any given odor is poor. The concentration of an odor-producing substance must be changed by about 30% before a difference can be detected. The comparable visual discrimination threshold is a 1% change in light intensity. The direction from which a smell comes may be indicated by the slight difference in the time of arrival of odoriferous molecules in the two nostrils. Odor-producing molecules are generally small, containing from 3 or 4 to 18-20 carbon atoms, and molecules with the same number of carbon atoms but different structural configurations have different odors. Relatively high water and lipid solubility are characteristic of substances with strong odors. Signal Transduction There are many different odorant receptors, and a second part is that the olfactory glomeruli to which the receptor cells project function as feature detectors. In mice, there are about 1000 different odorant receptors, and the number in humans is comparable. Since there are only 50,000-100,000 genes in the human genome, this means that up to 1% of the genome is devoted to making odorant receptors. All the odorant receptors are coupled to heterotrimeric G proteins. Some act via adenylyl cyclase and cAMP, and others act via phospholipase C and the products of phosphatidylinositol hydrolysis. Most of them open cation channels, causing an inward-directed Ca2+ current. Although there are 1000 different receptors, humans can discriminate ten times this many different odors. Consequently, an additional mechanism must be involved. It appears that this is provided by the olfactory glomeruli acting as feature detectors that project different patterns to the cortex. Each glomerulus receives input from only one type of olfactory receptor
However, each olfactory receptor responds to multiple odorants, and a single odorant can be recognized by multiple receptor types. Therefore, each odorant produces a different pattern of activated glomeruli, and the brain recognizes odors by the pattern of glomerular activation. In the olfactory glomeruli, there is lateral inhibition mediated by periglomerular cells and granule cells. This sharpens and focuses olfactory signals. In addition, the extracellular field potential in each glomerulus oscillates, and the granule cells appear to regulate the frequency of the oscillation. Olfactory stimuli appear to be terminated by protein kinases and enzymes that catalyze covalent modification of odor-producing molecules.
Odorant-Binding Proteins In contrast to the low threshold for olfactory stimulation when the olfactory mucous membrane is intact, single olfactory receptors that have been patch-clamped have a relatively high threshold and a long latency. In addition, lipophilic odor-producing molecules must traverse the hydrophilic mucus in the nose to reach the receptors. These facts led to the suggestion that the olfactory mucus might contain one or more odorant-binding proteins (OBP) that concentrate the odorants and transfer them to the receptors. An 18-kDa OBP that is unique to the nasal cavity has been isolated, and other related proteins probably exist. The protein has considerable homology to other proteins in the body that are known to be carriers for small lipophilic molecules. A similar binding protein appears to be associated with taste Vomeronasal Organ In rodents and various other mammals, the nasal cavity contains another patch of olfactory mucous membrane located along the nasal septum in a well-developed vomeronasal organ. This structure is concerned with the perception of odors that act as pheromone. Its receptors project to the accessory olfactory bulb and from there primarily to areas in the amygdala and hypothalamus that are concerned with reproduction and ingestive behavior. Vomeronasal input has major effects on these functions. The vomeronasal organ has about 30 serpentine odorant receptors that differ quite markedly in structure from those in the rest of the olfactory epithelium. The organ is not well developed in humans, but there is an anatomically separate and biochemically unique area of olfactory mucous membrane in a pit in the anterior third of the nasal septum which appears to be a homologous structure. The sense of smell is said to be more acute in women than in men, and in women it is most acute at the time of ovulation. Smell and, to a lesser extent, taste have a unique ability to trigger long-term memories, Sniffing The portion of the nasal cavity containing the olfactory receptors is poorly ventilated. Most of the air normally moves smoothly over the turbinates with each respiratory cycle, although eddy currents pass some air over the olfactory mucous membrane. These eddy currents are probably set up by convection as cool air strikes the warm mucosal surfaces. The amount of air reaching this region is greatly increased by sniffing, an action that includes contraction of the lower part of the nares on the septum to help deflect the airstream upward. Sniffing is a semi-reflex response that usually occurs when a new odor attracts attention.
Role of Pain Fibers in the Nose Naked endings of many trigeminal pain fibers are found in the olfactory mucous membrane. They are stimulated by irritating substances, and an irritative, trigeminally mediated component is part of the characteristic "odor" of such substances as peppermint, menthol, and chlorine. These endings are also responsible for initiating sneezing, lacrimation, respiratory inhibition, and other reflex responses to nasal irritants. Adaptation It is common knowledge that when one is continuously exposed to even the most disagreeable odor, perception of the odor decreases and eventually ceases. This sometimes beneficent phenomenon is due to the fairly rapid adaptation, or desensitization, that occurs in the olfactory system. It is specific for the particular odor being smelled, and the threshold for other odors is unchanged. It is at least partly central in origin and is apparently associated with hyperpolarization of cortical neurons. Abnormalities Abnormalities of olfaction include anosmia (absence of the sense of smell), hyposmia (diminished olfactory sensitivity), and dysosmia (distorted sense of smell). Several dozen different anosmias have been detected in humans. They are presumably due in each case to absence or disrupted function of one of the many members of the odorant receptor family. Olfactory thresholds increase with advancing age, and more than 75% of humans over the age of 80 have an impaired ability to identify smells.
RECEPTOR ORGANS & PATHWAYS Taste Buds The taste buds, the sense organs for taste, are ovoid bodies measuring 50-70 um. Each taste bud is made up of four types of cells basal cells; type 1 and 2 cells, which are sustentacular cells; and type 3 cells, which are the gustatory receptor cells that make synaptic connections to sensory nerve fibers. The type 1, 2, and 3 cells have microvilli, which project into the taste pore, an opening in the lingual epithelium. The necks of all these cells are connected to each other and to the surrounding epithelial cells by tight junctions, so that the only part of the gustatory receptor exposed to the fluids in the oral cavity is its apical crown of microvilli. Each taste bud is innervated by about 50 nerve fibers, and conversely, each nerve fiber receives input from an average of five taste buds. The basal cells arise from the epithelial cells surrounding the taste bud. They differentiate into new receptor cells, and the old receptor cells are continuously replaced with a half-time of about 10 days. If the sensory nerve is cut, the taste buds it innervates degenerate and eventually disappear. However, if the nerve regenerates, the cells in the neighborhood become organized into new taste buds, presumably as a result of some sort of chemical inductive effect from the regenerating fiber. In humans, the taste buds are located in the mucosa of the epiglottis, palate, and pharynx and in the walls of the fungiform and vallate papillae of the tongue. The fungiform papillae are rounded structures most numerous near the tip of the tongue; the vallate papillae are prominent structures arranged in a V on the back of the tongue. There are up to five taste buds per fungiform papilla, and they are usually located at the top of the papilla. The larger vallate papillae each contain up to 100 taste buds usually located along the sides of the papillae.
The small conical filiform papillae that cover the dorsum of the tongue do not usually contain taste buds. There are a total of about 10,000 taste buds.
Taste Pathways The sensory nerve fibers from the taste buds on the anterior two-thirds of the tongue travel in the chorda tympani branch of the facial nerve, and those from the posterior third of the tongue reach the brain stem via the glossopharyngeal nerve. The fibers from areas other than the tongue reach the brain stem via the vagus nerve. On each side, the myelinated but relatively slowly conducting taste fibers in these three nerves unite in the nucleus of the tractus solitarius in the medulla oblongata. There they synapse on second-order neurons, the axons of which cross the midline and join the medial lemniscus, ending with the fibers for touch, pain, and temperature sensibility in the specific sensory relay nuclei of the thalamus. Impulses are relayed from there to the taste projection area in the cerebral cortex at the foot of the postcentral gyrus. Taste does not have a separate cortical projection area but is represented in the portion of the postcentral gyrus that subserves cutaneous sensation from the face. Basic Taste Modalities In humans there are four established basic tastes: sweet, sour, bitter, and salt. There is considerable overlap, but bitter substances are tasted mostly on the back of the tongue, sour along the edges, sweet at the tip and salt on the dorsum anteriorly Sour and bitter substances are also tasted on the palate along with some sensitivity to sweet and salt. All four modalities can be sensed on the pharynx and epiglottis. Some taste cells respond best to bitter stimuli whereas others respond best to salt, sweet, or sour stimuli. Some respond to more than one modality, and some to all four. An additional taste modality named umami has been postulated to exist. It is said to mediate the taste of glutamate and the monosodium glutamate used extensively in Asian cooking. A variant metabotropic glutamate receptor, a truncated mGluR4, is said to be the receptor for this taste. Substances Evoking Primary Taste Sensations Acids taste sour. The H+ cation, rather than the associated anion, stimulates the receptors. For any given acid, sourness is generally proportionate to the H+ concentration, but organic acids are often more sour for a given H+ concentration than are mineral acids. This is probably because they penetrate cells more rapidly than the mineral acids do. A salty taste is produced by Na+. Some organic compounds also taste salty; for example, the dipeptides lysyltaurine and ornithyltaurine taste salty, and on a weight basis, lysyltaurine is more potent than NaCl. Most sweet substances are organic. Sucrose, maltose, lactose, and glucose are the most familiar examples, but polysaccharides, glycerol, some of the alcohols and ketones, and a number of compounds with no apparent relation to any of these, such as chloroform, beryllium salts, and various amides of aspartic acid, also taste sweet. Two proteins isolated from African berries, thaumatin and monellin, are 100,000 times as sweet as sucrose on a molar basis. The structures of these two proteins are very different, yet antibodies to one cross-react with antibodies to the other. Therefore, they must have some sort of common three-dimensional structure, and it may be this common feature that binds to the sweetness receptor. The substance usually used to test the bitter taste is quinine sulfate. This compound can be detected in a concentration of 8 umol/L; although the threshold for strychnine hydrochloride is
even lower. The taste is due to the cation. Thus, there is no apparent common feature of the molecular structure of substances that taste bitter. Taste Thresholds & Intensity Discriminations
- The ability of humans to discriminate differences in the intensity of tastes, like intensity
discrimination in olfaction, is relatively crude. A 30% change in the concentration of the substance being tasted is necessary before an intensity difference can be detected. The threshold concentrations of substances to which the taste buds respond vary with the particular substance
Receptor Stimulation The gustatory receptor cells are chemoreceptors that respond to substances dissolved in the oral fluids bathing them. These substances act on the exposed microvilli in the taste pore to evoke generator potentials in the receptor cells, which generate action potentials in the sensory neurons. The way the molecules in solution produce generator potentials varies from one gustatory modality to another. A protein called mammalian degenerin-1 may be the sour receptor. Alternatively, acids may depolarize sour receptor cells by activating H+-gated cation channels. Na+ depolarizes salt receptor cells via a Na+ channel that is related to the epithelial sodium channel (ENaC). ENaC is inhibited by amiloride, and direct application of this diuretic to the tongue in humans abolishes the ability to taste salt. Substances that taste sweet activate adenylyl cyclase via a heterotrimeric G protein, and the resulting increase in intracellular cAMP reduces K+ conductance by phosphorylating K+ channels in the basolateral membranes of the taste cells, producing depolarization. Substances that taste bitter reduce cAMP in their target taste cells, probably via a heterotrimeric G protein, and increase IP3 and DAG. A novel G protein named a-gusducin has been cloned and shown to occur in bitter taste cells. It activates phosphodiesterase, causing a decrease in intracellular cAMP. However, a-gusducin is also found in sweet taste cells, in which stimulation is associated with an increase intracellular cAMP. Thus, the exact role of a-gusducin remains unsettled. A protein that binds taste-producing molecules has been cloned. It is produced by Ebner's glands, glands that secrete mucus into the cleft around vallate papillae, and probably has a concentrating and transport function similar to that of the OBP described in the section on olfaction. Flavor The almost infinite varieties of tastes so dear to the gourmet are mostly synthesized from the four basic taste components. In some cases, a desirable taste includes an element of pain stimulation (e.g., "hot" sauces). In addition, smell plays an important role in the overall sensation produced by food, and the consistency (or texture) and temperature of foods also contributes to their "flavor." Variation & After-Effects There is considerable variation in the distribution of the four basic taste buds in various species and, within a given species, from individual to individual. In humans, there is an interesting variation in ability to taste phenylthiocarbamide (PTC).
In dilute solution, PTC tastes bitter to about 70% of the Caucasian population but is tasteless to the other 30%. Inability to taste PTC is inherited as an autosomal recessive trait. Testing for this trait is of considerable value in studies of human genetics. Taste exhibits after-reactions and contrast phenomena that are similar in some ways to visual after- images and contrasts. Some of these are chemical "tricks," but others may be true central phenomena. A taste modifier protein, miraculin, has been discovered in a plant. When applied to the tongue, this protein makes acids taste sweet. Animals, including humans, form particularly strong aversions to novel foods if eating the food is followed by illness. The survival value of such aversions is apparent in terms of avoiding poisons.
Abnormalities Abnormalities of taste include ageusia (absence of the sense of taste), hypogeusia (diminished taste sensitivity), and dysgeusia (disturbed sense of taste). Many different diseases can produce hypogeusia. In addition, drugs such as captopril and penicillamine, which contain sulfhydryl groups, cause temporary loss of taste sensation. The reason for this effect of sulfhydryl compounds is not known.
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