The Tongue

Roll no 72
Abdullah Shafique

GROSS
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.
 Nerve
Muscle Origin Insertion Supply Action
Intrinsic Muscles
Longitudinal Median septum Mucous Hypoglossal Alters shape of tongue
and submucosa membrane nerve
Transverse
Vertical
Extrinsic Muscles
GenioglossusSuperior genial Blends with other Hypoglossal Protrudes apex of tongue
spine of mandible muscles of nerve through mouth
tongue
Hyoglossus Body and greater Blends with other Hypoglossal Depresses tongue
cornu of hyoid muscles of nerve
bone tongue
Styloglossus Styloid process of Blends with other Hypoglossal Draws tongue upward and
temporal bone muscles of nerve backward
tongue
PalatoglossusPalatine Side of tongue Pharyngeal Pulls roots of tongue upward
aponeurosis plexus 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.
Development

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.
Histology

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.
Physiology

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.