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GUSTATORY SENSE Anatomic and Physiologic Considerations
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The sensory receptors for taste (taste buds) are distributed over the surface of the tongue and, in smaller numbers, over the soft palate, pharynx, larynx, and esophagus Mainly they are located in the epithelium along the lateral surfaces of the circumvallate and foliate papillae and to a lesser extent on the surface of the fungiform papillae The taste buds are round or oval structures, each composed of up to 200 vertically oriented receptor cells arranged like the staves of a barrel The super cial portion of the bud is marked by a small opening, the taste pore or pit, which opens onto the mucosal surface The tips of the sensory cells project through the pore as a number of liform microvilli ( taste hairs ) Fine, unmyelinated sensory bers penetrate the base of the taste bud and synapse directly with the sensory taste cells, which have no axons The taste receptors are activated by chemical substances in solution and transmit their activity along the sensory nerves to the brainstem There are four primary and readily tested taste sensations that have been long known: salty, sweet, bitter, and sour; recently a fth, umani the taste of glutamate, aspartate, and certain ribonucleotides has been added The full range of taste sensations is much broader, consisting of combinations of these elementary gustatory sensations Older notions of a tongue map, which implied the existence of speci c areas subserving one or another taste, are incorrect Any one taste bud is capable of responding to a number of sapid substances, but it is always preferentially sensitive to one type of stimulus In other words, the receptors are only relatively speci c The sensitivity of these receptors is remarkable: as little as 005 mg/dL of quinine sulfate will arouse a bitter taste when applied to the base of the tongue In recent years, a G-protein transduction system (gustaductin), similar to the one for olfaction, has been found to be operative in signaling taste sensations in the tongue receptors A discussion of this system can be found in the commentary by Brand cited in the References The receptor cells of the taste buds have a brief life cycle
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(about 10 days), being replaced constantly by mitotic division of adjacent basal epithelial cells The number of taste buds, not large to begin with, is gradually reduced with age; also, changes occur in the taste cell membranes, with impaired function of ion channels and receptors (Mistretta) Both gustatory (and olfactory) acuity diminish (everything begins to taste and smell the same) According to Schiffman, taste thresholds for salt, sweeteners, and amino acids are 2 to 2 1/2 times higher in the elderly than in the young The reduction in the acuity of taste and smell with aging may lead to a distortion of food habits (eg, excessive use of salt and other condiments) and contribute to the anorexia and weight loss of elderly persons Richter has explored the biologic role of taste in normal nutrition Animals made de cient in sodium, calcium, certain vitamins, proteins, etc, will automatically select the correct foods, on the basis of their taste, to compensate for their de ciency Interesting genetic polymorphisms in the receptor for sweet substances in rats have been found to underlie differences in the proclivity to ingest sweet substances, and a similar system has been proposed in humans (Chaudhari and Kinnamon) Neural Innervation of Tongue Regions Sensory impulses for taste arise from several sites in the oropharynx and are transmitted to the medulla via several cranial nerves (V, VII, IX, and X) The main pathway arises on the anterior two-thirds of the tongue; these taste bers rst run in the lingual nerve [a major branch of the mandibular-trigeminal ( fth) cranial nerve] After coursing within the lingual nerve for a short distance, the taste bers diverge to enter the chorda tympani (a branch of the seventh nerve); thence they pass through the pars intermedia and geniculate ganglion of the seventh nerve to the rostral part of the nucleus of the tractus solitarius in the medulla, where all taste afferents converge (see below) Fibers from the palatal taste buds pass through the pterygopalatine ganglion and greater super cial petrosal nerve, join the facial nerve at the level of the geniculate ganglion, and proceed to the nucleus of the solitary tract (see Fig 47-3) Possibly, some taste bers from the tongue may also reach the brainstem via the mandibular division of the trigeminal nerve The presence of this alternative pathway probably accounts for reported instances of unilateral taste loss that have followed section of the root of the trigeminal nerve and instances in which no loss of taste has occurred with section of the chorda tympani From the posterior third of the tongue, soft palate, and palatal arches, the sensory taste bers are conveyed via the glossopharyngeal nerve and ganglion nodosum to the nucleus of the tractus solitarius Taste bers from the extreme dorsal part of the tongue and the few that arise from taste buds on the pharynx and larynx run in the vagus nerve Rostral and lateral parts of the nucleus tractus solitarius, which receive the special afferent (taste) bers from the facial and glossopharyngeal nerves, constitute the gustatory nucleus Probably both sides of the tongue are represented in this nucleus The second sensory neuron for taste has been dif cult to track Neurons of the nucleus solitarius project to adjacent nuclei (eg, dorsal motor nucleus of the vagus, ambiguus, salivatorius superior and inferior, trigeminal, and facial nerves), which serve viscerovisceral and viscerosomatic re ex functions, but those concerned with the conscious recognition of taste are believed to form an ascending pathway to a pontine parabrachial nucleus From the latter, two ascending pathways have been traced (in animals) One is the solitariothalamic lemniscus to the ventroposteromedial nu-
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cleus of the thalamus A second passes to the ventral parts of the forebrain, to parts of the hypothalamus (which probably in uences autonomic function), and to other basal forebrain limbic areas in or near the uncus of the temporal lobe Other ascending bers lie near the medial lemniscus and are both crossed and uncrossed Experiments in animals indicate that taste impulses from the thalamus project to the tongue-face area of the postrolandic sensory cortex This is probably the end station of gustatory projections in humans as well, insofar as gustatory hallucinations have been produced by electrical stimulation of the parietal and/or rolandic opercula (Hausser-Hauw and Bancaud) Pen eld and Faulk evoked distinct taste sensations by stimulating the anterior insula
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