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Sense Organs / 235

               called taste cells, are arranged in a group   part of the tongue is conveyed by fibers
                                                        of the glossopharyngeal nerve. Somatic
               with supporting cells, a cluster that consti­
  VetBooks.ir  tutes the taste bud (Fig. 12‐4). Taste buds   sensations (heat, cold, touch, pain) from
                                                        the tongue are conveyed by branches of the
               are not distributed evenly on the surface of
               the tongue; they are confined to specific   trigeminal and glossopharyngeal nerves.
               forms of papillae (tiny projections), the   Taste buds on areas other than the tongue
               vallate, foliate, and fungiform papillae,   (e.g., soft palate) are innervated by the
               which are found on the tongue (the greatest   vagus nerve.
               number), soft palate, parts of the pharynx,   Traditionally, four basic taste sensations
               and epiglottis of the larynx.            had been identified. These are  sweet,
                  Sensory nerve fibers subserving taste   salt,  bitter, and  sour. Individual taste
               are  distributed  to  the  rostral  portions  of   cells have membrane receptor physiology
               the tongue by a branch of the facial nerve,   designed to detect the chemical substances
               the chorda tympani. Taste in the caudal   associated with these tastes. The more
                                                        complex sensory experiences that we
                (A)                                     normally associate with taste (for example,
                                                        the flavors that we detect when we distin­
                                                        guish between an apple and a carrot) are
                                                        created primarily from stimulation of
                                                        olfactory (smell) receptors in combination
                                                        with the basic taste modalities. For them to
                                                        stimulate taste cells, chemicals must be in
                                                        solution. Dissolution of substances so
                                                        that  they  can be tasted is an important
                                                        function of saliva.
                                                           In addition to the four basic tastes, a fifth
                                                        taste modality with its own specific taste
                                                        cell receptor is also recognized. This taste,
                                                        experienced when the amino acid gluta­
                                                        mate is present along with sodium, imparts
                (B)                                     a  savory quality to  foodstuffs.  Because
                                                        this taste modality was first described by
                                                        Japanese researchers, it is known by the
                                                        Japanese word for savory or delicious:
                                                        umami. It is the presence of the umami
                                                        taste receptor that  explains the  enhanced
                                                        flavor of foods when monosodium gluta­
                                                        mate (MSG) is added.



                                                        Olfaction

                                                        Olfaction is the sense of smell. Olfactory
                                                        sensory neurons are scattered among
               Figure  12-4.  Photomicrograph of lingual   supporting cells throughout the olfactory
               papillae. (A) 10 ×, box indicates single taste bud
               on lateral aspect of papilla. (B) 40 × of several   mucosa in the dorsocaudal part of the nasal
               taste buds; arrow indicates opening (taste pore)   cavity (Fig. 12‐5). The apex of each olfac­
               into bud which allows dissolved substances to   tory neuron bears a single dendrite with a
               contact taste cells. Source: micrograph courtesy of   tuft of several fine hair‐like projections,
               Gretchen Delcambre, Colorado State University,   which bear the chemical receptors for the
               Fort Collins, Colorado, USA.             sense of smell. An axon from each olfactory
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