The Tongue Roll no 72 Abdullah Shafique

The tongue is a mass of striated muscle covered with mucous membrane. The muscles attach the tongue to the styloid process and the soft palate above and to the mandible and the hyoid bone below. The tongue is divided into right and left halves by a median fibrous septum.

Mucous Membrane of the Tongue The mucous membrane of the upper surface of the tongue can be divided into anterior and posterior parts by a V-shaped sulcus, the sulcus terminalis (Fig. 1). The sulcus serves to divide the tongue into the anterior two thirds, or oral part, and the posterior third, or pharyngeal part. The foramen cecum is an embryologic remnant and marks the site of the upper end of the thyroglossal duct.

Figure 1. Dorsal surface of the tongue showing the valleculae, the epiglottis, and the entrance into the piriform fossa on each side (arrows). Three types of papillae are present on the upper surface of the anterior two thirds of the tongue: the filiform papillae, the fungiform papillae, and the vallate papillae. The mucous membrane covering the posterior third of the tongue is devoid of papillae but has an irregular surface caused by the presence of underlying lymph nodules, the lingual tonsil. The mucous membrane on the inferior surface of the tongue is reflected from the tongue to the floor of the mouth. In the midline anteriorly, the undersurface of the tongue is connected to the floor of the mouth by a fold of mucous membrane, the frenulum of the tongue. On the lateral side of the frenulum, the deep lingual vein can be seen through the mucous membrane. Lateral to the lingual vein, the mucous membrane forms a fringed fold called the plica fimbriata. Muscles of the Tongue The muscles of the tongue are divided into two types: intrinsic and extrinsic.

Intrinsic Muscles These muscles are confined to the tongue and are not attached to bone. They consist of longitudinal, transverse, and vertical fibers.
• •

Nerve supply: Hypoglossal nerve Action: Alter the shape of the tongue

Extrinsic Muscles These muscles are attached to bones and the soft palate. They are the genioglossus, the hyoglossus, the styloglossus, and the palatoglossus.

Nerve supply: Hypoglossal nerve

The origin, insertion, nerve supply, and action of the tongue muscles are summarized in the above Table.

Muscle Origin Intrinsic Muscles Longitudinal Median septum and submucosa Transverse Vertical Extrinsic Muscles GenioglossusSuperior genial spine of mandible

Insertion Mucous membrane

Nerve Supply


Hypoglossal Alters shape of tongue nerve

Blends with other Hypoglossal muscles of nerve tongue Hyoglossus Body and greater Blends with other Hypoglossal cornu of hyoid muscles of nerve bone tongue Styloglossus Styloid process of Blends with other Hypoglossal temporal bone muscles of nerve tongue PalatoglossusPalatine Side of tongue Pharyngeal aponeurosis plexus

Protrudes apex of tongue through mouth Depresses tongue Draws tongue upward and backward Pulls roots of tongue upward and backward, narrows oropharyngeal isthmus

Blood Supply The lingual artery, the tonsillar branch of the facial artery, and the ascending pharyngeal artery supply the tongue. The veins drain into the internal jugular vein. Lymph Drainage
• • •

Tip: Submental lymph nodes Sides of the anterior two thirds: Submandibular and deep cervical lymph nodes Posterior third: Deep cervical lymph nodes

Sensory Innervation
• •

Anterior two thirds: Lingual nerve branch of mandibular division of trigeminal nerve (general sensation) and chorda tympani branch of the facial nerve (taste) Posterior third: Glossopharyngeal nerve (general sensation and taste)

Movements of the Tongue
• • • • •

Protrusion: The genioglossus muscles on both sides acting together (Figure 2) Retraction: Styloglossus and hyoglossus muscles on both sides acting together Depression: Hyoglossus muscles on both sides acting together Retraction and elevation of the posterior third: Styloglossus and palatoglossus muscles on both sides acting together Shape changes: Intrinsic muscles

Figure 2. Diagrammatic representation of the action of the right and left genioglossus muscles of the tongue. A. The right and left muscles contract equally together and as a result (B) the tip of the tongue is protruded in the midline. C. The right hypoglossal nerve (which innervates the genioglossus muscle and the intrinsic tongue muscles on the same side) is cut and as a result the right side of the tongue is atrophied and wrinkled. D. When the patient is asked to protrude the tongue, the tip points to the side of the nerve lesion. E. The origin and insertion and direction of pull of the genioglossus muscle.

A small median elevation, the tuberculum impar or median tongue bud, appears in the floor of the pharynx before the pharyngeal arches meet ventrally, and it subsequently becomes incorporated in the anterior part of the tongue. A little later two oval mandibular or lingual swellings appear on the inner aspect of the mandibular processes. They meet each other in front, and caudally they converge on the tuberculum impar, with which they fuse. A sulcus forms along the ventral and lateral margins of this elevation and deepens, internal to the future alveolar process of the mandible, to form the linguogingival groove, while the elevation constitutes the anterior or buccal (presulcal) part of the tongue. Caudal to the tuberculum impar, a second median elevation, the hypobranchial eminence (copula of His), forms in the floor of the pharynx, and the ventral ends of the fourth, third and, later, second, pharyngeal arches converge into it. A transverse groove separates its caudal part to delineate the epiglottis. Ventrally it approaches the presulcal tongue rudiment, and spreads in the form of a V, to form the posterior or pharyngeal part of the tongue. During this sequence of events, the third arch elements grow over and bury the elements of the second arch, thereby excluding it from the tongue. The mucous membrane of the pharyngeal part of the tongue therefore receives its sensory supply from the glossopharyngeal nerve, which is the nerve of the third arch. In the adult, the union of the anterior and posterior parts of the tongue approximately corresponds to the angulated sulcus terminalis, which has its apex at the foramen caecum, a blind depression produced at the time of fusion of the constituent parts of the tongue, but also marking the site of ingrowth of the median rudiment of the thyroid gland. The tongue initially consists of a mass of mesenchyme covered on its surface by first arch ectoderm and third arch endoderm. During the second month it is invaded by occipital myotomes which migrate from the lateral aspects of the myelencephalon. They pass ventrally round the pharynx to reach its floor accompanied by the hypoglossal nerve. The composite character of the tongue is revealed by its sensory innervation. The anterior, buccal part is innervated by the lingual nerve, derived from the post-trematic nerve of the first arch (mandibular nerve) and by the chorda tympani, which is often held to be the pretrematic nerve to the first arch. The posterior, pharyngeal part of the tongue is innervated by the glossopharyngeal, the nerve of the third arch, and the root of the tongue, near the epiglottis, is innervated by the vagus. The sulcus terminalis cannot be distinguished earlier than the 52 mm stage according to some observers. The vallate papillae appear at about the same time, and increase in number until the 170 mm stage. Serial reconstructions suggest that the territory of the glossopharyngeal nerve extends considerably beyond these papillae. Lymphoid tissue similar to the palatine tonsils usually develops on the surface of the posterior part of the tongue, and is called the lingual tonsil.

In the neonate, the tongue is short and broad, and its entire surface lies within the oral cavity. The posterior third of the tongue descends into the neck during the first postnatal year, and by the fourth or fifth year the tongue forms part of the anterior wall of the pharynx.


The dorsal surface of the tongue is divided by the sulcus terminalis into an oral part, the anterior two-thirds, and a pharyngeal part, the posterior one-third. The dorsal surface of the oral part has a characteristic appearance due to the presence of a large number of small projections, the lingual papillae. The epithelium of the pharyngeal part forms a somewhat irregular surface which covers the lingual tonsils.

The lingual papillae consist of a connective tissue core covered with a stratified squamous epithelium. On the basis of their appearance four types of papillae can be distinguished filiform, fungiform, circumvallate and foliate papillae.

Filiform papillae Filiform papillae are minute, conical or cylindrical projections which cover most of the presulcal dorsal area, and are arranged in diagonal rows that extend anterolaterally, parallel with the sulcus terminalis, except at the lingual apex where they are transverse. They have irregular cores of connective tissue and their epithelium, which is keratinized, may split into whitish fine secondary processes. They appear to function to increase the friction between the tongue and food, and facilitate the movement of particles by the tongue within the oral cavity.

Fungiform papillae Fungiform papillae occur mainly on the lingual margin but also irregularly on the dorsal surface, where they may occasionally be numerous. They differ from filiform papillae because they are larger, rounded and deep red in colour, this last reflecting their thin, non-keratinized epithelium and highly vascular connective tissue core. Each usually bears one or more taste buds on its apical surface.

Dorsal surface of the anterior tongue showing non-keratinized fungiform (left) and two keratinized filiform papillae (centre and right) with non-keratinized regions between. (By permission from Young B, Heath JW 2000 Wheater's Functional Histology. Edinburgh: Churchill Livingstone.)

Foliate papillae Foliate papillae lie bilaterally in two zones at the sides of the tongue near the sulcus terminalis, each formed by a series of red, leaf-like mucosal ridges, covered by a non-keratinized epithelium. They bear numerous taste buds.

Circumvallate papillae Circumvallate papillae are large cylindrical structures, varying in number from 8 to 12, which form a V-shaped row immediately in front of the sulcus terminalis on the dorsal surface of the tongue. Each papilla, 1-2 mm in diameter, is surrounded by a slight circular mucosal elevation (vallum or wall) which is separated from the papilla by a circular sulcus. The papilla is narrower at its base than its apex and the entire structure is generally covered with non-keratinized stratified squamous epithelium. Numerous taste buds are scattered in both walls of the sulcus, and small serous glands (of von Ebner) open into the sulcal base.

Section through a circumvallate papilla. Serous glands (of von Ebner) empty via ducts into the base of the trench and numerous taste buds are contained within the stratified epithelium of the papillary wall (pale structures on the inner wall of the cleft, left side). (By permission from Young B, Heath JW 2000 Wheater's Functional Histology. Edinburgh: Churchill Livingstone.)

The epithelium of the dorsal surface of the tongue rests on a fairly dense layer of connective tissue, which connects the epithelium firmly with the underlying muscular and connective tissues. The muscles of the tongue (skeletal muscle) are organized into strands oriented more or less perpendicular to each other. Their actions provide the tongue with the necessary motility to participate in the formation of speech and to aid in the initial processing of foods. Control of the movement of the tongue muscles and the collection of sensory information necessitate a profuse innervation of the tongue in which a number of the cranial nerves participate (V, trigeminal nerve sensory - anterior two-thirds; VII, facial nerve - taste; IX, glossopharyngeal nerve - sensory/taste - posterior one-third; XII, hypoglossal nerve - motor).

Taste buds
Taste buds are microscopic barrel-shaped epithelial structures which contain chemosensory cells in synaptic contact with the terminals of gustatory nerves. They are numerous on all types of lingual papillae (except filiform papillae) particularly on their lateral aspects. Taste buds are not restricted to the papillae, and are scattered over almost the entire dorsal and lateral surfaces of the tongue and, rarely, on the epiglottis and lingual aspect of the soft palate. Each taste bud is linked by synapses at its base to one of three cranial nerves which carry taste, i.e. the facial, glossopharyngeal or vagus. They share some physiological features with neurones, for example action potential generation and synaptic transmission, and are therefore often referred to as paraneurones.

Figure 33.12 Circumvallate papilla. A, Scanning electron micrograph showing a circumvallate papilla surrounded by a trench. B, Section of circumvallate papilla showing pale barrel-shaped taste buds (B) in its walls. P, apical pore. (A, by kind permission from S Franey and by permission from Berkovitz BKB, Holland GR, Moxham BJ 2002 Oral Anatomy, Embryology and Histology, 3rd edn. Edinburgh: Mosby; B, by permission from Dr JB Kerr, Monash University, from Kerr JB 1999 Atlas of Functional Histology. London: Mosby.) There is considerable individual variation in the distribution of taste buds in humans. They are

most abundant on the posterior parts of the tongue, especially around the walls of the circumvallate papillae and their surrounding sulci, where there is an average of c.250 taste buds for each of the 8-12 papillae. Over 1000 taste buds are distributed over the sides of the tongue, particularly over the more posterior folds of the two foliate papillae, whereas they are rare, and sometimes even absent, on fungiform papillae (c.3 per papilla). Taste buds have been described on the fetal epiglottis and soft palate but most disappear from these sites during postnatal development. Each taste bud is a barrel-shaped cluster of 50-150 fusiform cells which lies within an oval cavity in the epithelium and converges apically on a gustatory pore, a 2 μm wide opening on the mucosal surface. The whole structure is about 70 μm in height by 40 μm across and is separated by a basal lamina from the underlying lamina propria. A small fasciculus of afferent nerve fibres penetrates the basal lamina and spirals around the sensory cells. Chemical substances dissolved in the oral saliva diffuse through the gustatory pores of the taste buds to reach the taste receptor cell membranes, where they cause membrane depolarization. Individual nerve fibres branch to give a complex distribution of taste bud innervation. Each fibre may have many terminals, which may spread to innervate widely separated taste buds or may innervate more than one sensory cell in each bud. Conversely, individual buds may receive the terminals of several different nerve fibres. These convergent and divergent patterns of innervation may be of considerable functional importance. The gustatory nerve for the anterior part of the tongue, excluding the circumvallate papillae, is the chorda tympani, which travels via the lingual nerve. In most individuals, taste fibres run in the chorda tympani to cell bodies in the facial ganglion, but occasionally they diverge to the otic ganglion, which they reach via the greater petrosal nerve. Taste buds in the inferior surface of the soft palate are supplied mainly by the facial nerve, through the greater petrosal nerve, pterygopalatine ganglion and lesser palatine nerve: they may also be supplied by the glossopharyngeal nerve. Taste buds in the circumvallate papillae, postsulcal part of the tongue and in the palatoglossal arches and the oropharynx are innervated by the glossopharyngeal nerve, and those in the extreme pharyngeal part of the tongue and epiglottis receive fibres from the internal laryngeal branch of the vagus. Each taste bud receives two distinct classes of fibre: one branches in the periphery of the bud to form a perigemmal plexus, the other forms an intragemmal plexus within the bud itself which innervates the bases of the receptor cells. The perigemmal fibres contain various neuropeptides including calcitonin gene-related peptide (CGRP) and substance P, and appear to represent free sensory endings. Intragemmal fibres branch within the taste bud and each forms a series of synapses.

Sense of Taste
Taste is mainly a function of the taste buds in the mouth, but it is common experience that one's sense of smell also contributes strongly to taste perception. In addition, the texture of food, as detected by tactual senses of the mouth, and the presence of substances in the food that stimulate pain endings, such as pepper, greatly alter the taste experience. The importance of taste lies in the fact that it allows a person to select food in accord with desires and often in accord with the body tissues' metabolic need for specific substances.

Primary Sensations of Taste

The identities of the specific chemicals that excite different taste receptors are not all known. Even so, psychophysiologic and neurophysiologic studies have identified at least 13 possible or probable chemical receptors in the taste cells, as follows: 2 sodium receptors, 2 potassium receptors, 1 chloride receptor, 1 adenosine receptor, 1 inosine receptor, 2 sweet receptors, 2 bitter receptors, 1 glutamate receptor, and 1 hydrogen ion receptor.

For practical analysis of taste, the aforementioned receptor capabilities have also been grouped into five general categories called the primary sensations of taste. They are sour, salty, sweet, bitter, and "umami."

A person can perceive hundreds of different tastes. They are all supposed to be combinations of the elementary taste sensations, just as all the colors we can see are combinations of the three primary colors, as described in Chapter 50.

Sour Taste. The sour taste is caused by acids, that is, by the hydrogen ion concentration, and the intensity of this taste sensation is approximately proportional to the logarithm of the hydrogen ion concentration. That is, the more acidic the food, the stronger the sour sensation becomes.

Salty Taste. The salty taste is elicited by ionized salts, mainly by the sodium ion concentration. The quality of the taste varies somewhat from one salt to another, because some salts elicit other taste sensations in addition to saltiness. The cations of the salts, especially sodium cations, are mainly responsible for the salty taste, but the anions also contribute to a lesser extent.

Sweet Taste. The sweet taste is not caused by any single class of chemicals. Some of the types of chemicals that cause this taste include sugars, glycols, alcohols, aldehydes, ketones, amides, esters, some amino acids, some small proteins, sulfonic acids, halogenated acids, and inorganic salts of lead and beryllium. Note specifically that most of the substances that cause a sweet taste are organic chemicals. It is especially interesting that slight changes in the chemical structure, such as addition of a simple radical, can often change the substance from sweet to bitter.

Bitter Taste. The bitter taste, like the sweet taste, is not caused by any single type of chemical agent. Here again, the substances that give the bitter taste are almost entirely organic substances. Two particular classes of substances are especially likely to cause bitter taste sensations: (1) long-chain organic substances that contain nitrogen, and (2) alkaloids. The alkaloids include many of the drugs used in medicines, such as quinine, caffeine, strychnine, and nicotine.

Some substances that at first taste sweet have a bitter aftertaste. This is true of saccharin, which makes this substance objectionable to some people.

The bitter taste, when it occurs in high intensity, usually causes the person or animal to reject the food. This is undoubtedly an important function of the bitter taste sensation, because many deadly toxins found in poisonous plants are alkaloids, and virtually all of these cause intensely bitter taste, usually followed by rejection of the food.

Taste Bud and Its Function

Figure. Taste bud.

The above shows a taste bud, which has a diameter of about 1/30 millimeter and a length of about 1/16 millimeter. The taste bud is composed of about 50 modified epithelial cells, some of which are supporting cells called sustentacular cells and others of which are taste cells. The taste cells are continually being replaced by mitotic division of surrounding epithelial cells, so that some taste cells are young cells. Others are mature cells that lie toward the center of the bud; these soon break up and dissolve. The life span of each taste cell is about 10 days in lower mammals but is unknown for humans.

The outer tips of the taste cells are arranged around a minute taste pore, shown inFigure. From the tip of each taste cell, several microvilli, or taste hairs, protrude outward into the taste pore to approach the cavity of the mouth. These microvilli provide the receptor surface for taste.

Interwoven around the bodies of the taste cells is a branching terminal network of taste nerve fibers that are stimulated by the taste receptor cells. Some of these fibers invaginate into folds of the taste cell membranes. Many vesicles form beneath the cell membrane near the fibers. It is believed that these vesicles contain a neurotransmitter substance that is released through the cell membrane to excite the nerve fiber endings in response to taste stimulation.

Location of the Taste Buds. The taste buds are found on three types of papillae of the tongue, as follows: (1) A large number of taste buds are on the walls of the troughs that surround the circumvallate papillae, which form a V line on the surface of the posterior tongue. (2) Moderate numbers of taste buds are on the fungiform papillae over the flat anterior surface of the tongue. (3) Moderate numbers are on the foliate papillae located in the folds along the lateral surfaces of the tongue. Additional taste buds are located on the palate, and a few are found on the tonsillar pillars, on the epiglottis, and even in the proximal esophagus. Adults have 3000 to 10,000 taste buds, and children have a few more. Beyond the age of 45 years, many taste buds degenerate, causing the taste sensation to become progressively less critical in old age.

Specificity of Taste Buds for a Primary Taste Stimulus. Microelectrode studies from single taste buds show that each taste bud usually responds mostly to one of the five primary taste stimuli when the taste substance is in low concentration. But at high concentration, most buds can be excited by two or more of the primary taste stimuli, as well as by a few other taste stimuli that do not fit into the "primary" categories.

Mechanism of Stimulation of Taste Buds Receptor Potential. The membrane of the taste cell, like that of most other sensory receptor cells, is negatively charged on the inside with respect to the outside. Application of a taste substance to the taste hairs causes partial loss of this negative potential-that is, the taste cell becomes depolarized. In most instances, the decrease in potential, within a wide range, is approximately proportional to the logarithm of concentration of the stimulating substance. This change in electrical potential in the taste cell is called the receptor potential for taste.

The mechanism by which most stimulating substances react with the taste villi to initiate the receptor potential is by binding of the taste chemical to a protein receptor molecule that lies on the outer surface of the taste receptor cell near to or protruding through a villus membrane. This, in turn, opens ion channels, which allows positively charged sodium ions or hydrogen ions to enter and depolarize the normal negativity of the cell. Then the taste chemical itself is gradually washed away from the taste villus by the saliva, which removes the stimulus.

The type of receptor protein in each taste villus determines the type of taste that will be perceived. For sodium ions and hydrogen ions, which elicit salty and sour taste sensations, respectively, the receptor proteins open specific ion channels in the apical membranes of the taste cells, thereby activating the receptors. However, for the sweet and bitter taste sensations, the portions of the receptor protein

molecules that protrude through the apical membranes activate second-messenger transmitter substances inside the taste cells, and these second messengers cause intracellular chemical changes that elicit the taste signals.

Generation of Nerve Impulses by the Taste Bud. On first application of the taste stimulus, the rate of discharge of the nerve fibers from taste buds rises to a peak in a small fraction of a second but then adapts within the next few seconds back to a lower, steady level as long as the taste stimulus remains. Thus, a strong immediate signal is transmitted by the taste nerve, and a weaker continuous signal is transmitted as long as the taste bud is exposed to the taste stimulus.

Transmission of Taste Signals into the Central Nervous System

Figure Transmission of taste signals into the central nervous system.

The above figure shows the neuronal pathways for transmission of taste signals from the tongue and pharyngeal region into the central nervous system. Taste impulses from the anterior two thirds of the tongue pass first into the lingual nerve, then through the chorda tympani into the facial nerve, and finally into the tractus solitarius in the brain stem. Taste sensations from the circumvallate papillae on the back of the tongue and from other posterior regions of the mouth and throat are transmitted through the glossopharyngeal nerve also into the tractus solitarius, but at a slightly more posterior level. Finally, a few taste signals are transmitted into the tractus solitarius from the base of the tongue and other parts of the

pharyngeal region by way of the vagus nerve.

All taste fibers synapse in the posterior brain stem in the nuclei of the tractus solitarius. These nuclei send second-order neurons to a small area of the ventral posterior medial nucleus of the thalamus, located slightly medial to the thalamic terminations of the facial regions of the dorsal column-medial lemniscal system. From the thalamus, third-order neurons are transmitted to the lower tip of the postcentral gyrus in the parietal cerebral cortex, where it curls deep into the sylvian fissure, and into the adjacent opercular insular area. This lies slightly lateral, ventral, and rostral to the area for tongue tactile signals in cerebral somatic area I. From this description of the taste pathways, it is evident that they closely parallel the somatosensory pathways from the tongue. Taste Reflexes Integrated in the Brain Stem. From the tractus solitarius, many taste signals are transmitted within the brain stem itself directly into the superior and inferior salivatory nuclei, and these areas transmit signals to the submandibular, sublingual, and parotid glands to help control the secretion of saliva during the ingestion and digestion of food. Adaptation of Taste. Everyone is familiar with the fact that taste sensations adapt rapidly, often almost completely within a minute or so of continuous stimulation. Yet, from electrophysiologic studies of taste nerve fibers, it is clear that adaptation of the taste buds themselves usually accounts for no more than about half of this. Therefore, the final extreme degree of adaptation that occurs in the sensation of taste almost certainly occurs in the central nervous system itself, although the mechanism and site of this are not known. At any rate, it is a mechanism different from that of most other sensory systems, which adapt almost entirely at the receptors.

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