ORAL BIOLOGY

TASTE

OVERVIEW
 RELEVANT ANATOMY OF THE TONGUE
Papillae
Innervation
Distribution of taste sensitivities on the tongue
 THE STRUCTURE OF TASTE BUDS
 SENSORY TRANSDUCTION
Type III cells - Salt, Acid
Type II cells - Sweet, Umami, Bitter
 RESPONSES OF TASTE AFFERENTS
 INFORMATION EXTRACTION
 CNS TASTE PATHWAYS
 NEUROTROPHISM AND TASTE BUDS

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INTRODUCTION
Taste and olfaction are examples of the specialised sensory systems that have evolved in
vertebrates for sensing the external environment.

Taste, or gustation, has often been considered to have limited value as a special sense because it
has little contribution to make in providing information about the distant environment. Taste
requires the direct application of a substance to the taste buds for it to be sensed, so in most
animals its value is assumed to be restricted to food identification and rejection. This is its main
importance in humans. Taste also initiates cephalic (neural) phase responses during ingestion
of food. The cephalic phase of nutrition refers to a set of food intake-associated autonomic and
endocrine responses upon stimulation of sensory pathways in the oropharynx. They include
salivation, production of hydrochloric acid, gastrin, digestive enzymes and immunoglobulins in
the stomach, pancreatic release of bicarbonate, insulin and glucagon and there is an anticipatory
increase in gastric motility.

It should also be noted that although specialised taste buds are found only in the mouth and
pharyngeal spaces in man, chemosensory cells possessing the same transduction apparatus as
found in taste cells are also found throughout the gastro-intestinal (GI) tract. This implies that the
GI tract has the ability to detect the same constituents of food that are detected in the mouth. This
system contributes to the coordination of enzyme release for digestion in response to specific
dietary components, and may also be involved in monitoring dietary content.

Flavour is the combined sensory experience of olfaction and gustation.

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TASTE

RELEVANT ANATOMY OF THE TONGUE

Papillae
The tongue carries four types of papilla, but not all types are associated with taste buds. The
situation is as summarised below.

Type of papillae Taste buds?

Filiform X
Fungiform 
Foliate 
Circumvallate 

Taste buds are also found in the epithelium of the soft palate, the oropharynx and around the
epiglottis

Innervation
The table summarises the anatomy however review the powerpoint slides used in the lecture.

Area of the tongue General sensation Gustation

(touch, temperature, (taste)
nociception)
Anterior 2/3 of tongue Cranial Nerve (CN) V CN VII
(via chorda tympani
branch, however note
lingual nerve of CN V at
beginning)
Posterior 1/3 of tongue CN IX CN IX
Root of tongue, epiglottis, CN IX and X CN X
oropharynx

Distribution of taste sensitivities on the tongue

It is frequently claimed that the tongue has distinct areas of sensitivity to the four basic modalities
of taste –sweet, salt, acid and bitter. This, however, is an oversimplification of the real picture.

The first tongue maps were described by D.P.Hänig in 1901, and these show relative sensitivities
to the different taste modalities, not absolute ones. He showed that all parts of the tongue are
sensitive to all four modalities, but there are differences in relative sensitivity as follows.

SWEET greatest sensitivity on the tip of the tongue

SALT greatest sensitivity on the antero-lateral aspects of the tongue
ACID greatest sensitivity on the postero-lateral aspects of the tongue
BITTER greatest sensitivity on the circumvallate papillae and the
posterior 1/3 of the tongue

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The picture has become more complex as it is now established that humans have a fifth taste
modality, which is referred to as “UMAMI” taste. The word “umami” comes from the Japanese
word for “delicious”, and this refers to the fact that stimulation of umami receptors seems to
enhance the pleasurable aspects of taste. The physiological agonist (excitant) of umami receptors
in taste buds is amino acids, and L-glutamate in particular. This is present in meat extracts,
certain cheeses and soy sauce. We know that humans have greater glutamate sensitivity at the
back of the tongue, and although there are slight differences across the anterior tongue these are
not significantly different in most people.
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THE STRUCTURE OF TASTE BUDS

[This section should be read in conjunction with the Figure 1 on the next page- an illustration of a
taste bud]
 All taste buds have the same basic structure. Each taste bud is embedded in the epithelium of
the oral mucosa, and is made up of 50 – 70 cells. These are arranged in much the same way
as the segments of an orange that has been peeled. The cells regenerate every 10 days.
 There is an apical pore that is open to the environment of the oral cavity. However, the nerve
supply enters the taste bud from the base. There are tight junctions between all the taste bud
cells at their apical ends, creating a waterproof seal.
 Electron microscopy studies have been used to describe the types of cells in the taste bud as
Types I, II, III, and IV.
 Taste cells originate from precursor cells resident in the surrounding epithelium.
 Type I cells (55-65% of cells). Thought to have glia-like properties. Elongated cells
extending through the full depth of taste bud, with about 40 microvilli at the apical end of the
cells. They have extended cytoplasmic lamellae that engulf other cells. May have a supportive
function, as they are wrapped around groups of other elongated taste cells. They have recently
been found to express amiloride sensitive Na+ channels i.e. they may respond to salty
stimuli (See following pages).
 Type II cells (20-30% of cells). Also known as receptor cells. Extend through full depth of
taste bud. They have synapse-like relations with nerve processes. They have G-protein
coupled receptors for sweet, bitter or umami compounds.
 Type III cells (10-15% of cells). Extend through the full depth of the taste bud. They have
no microvilli at the apical end. They are closely associated with nerve processes in the taste
bud and express synaptic proteins. Therefore also known as presynaptic cells. The cells
contain clusters of vesicles and there are areas of synapse-like specialisation with the nerve
processes. Individual nerve processes form synapses with more than one Type III cell in the
taste bud. There is substantial evidence that 5-hydroxytryptamine (5-HT)(serotonin) is the
transmitter at these synapses, but you should be aware that GABA and nor-adrenaline have
also been proposed as the transmitter.
 Type IV cells (basal cells). They do not extend processes into the taste pore. Their function is
unknown though they are likely to be undifferentiated or immature taste cells.
 Special sensory (taste or gustatory afferent) neurones penetrate the basal lamina to enter taste
buds. Their cell bodies are located in ganglia nestled against the brain (i.e. geniculate,
petrosal and nodose cranial ganglia). These sensory neurones intermingle with other nerves
under the epithelium including those that sense pain or thermal signals.

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Figure 1: Cell types in a taste bud

SENSORY TRANSDUCTION
Type II and Type III cells are involved in the transduction of gustatory stimuli, and they show
modality specificity as set out below. A particular Type II or Type III cell carries the
transduction mechanism for only one tastant. Thus Type III cells are either salt detecting or acid
detecting, but not both, and similarly for Type II cells.

Cell type Taste modalities

Type III Salt
Acid
Type II Sweet
Umami
Bitter

The transduction mechanisms for each of the five taste modalities will now be summarised.

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Salt
In humans, the evidence supports there being at least two possible mechanisms of salt
transduction. At the most basic level, Na+ can pass directly into the Type III cells through a Na+
permeable channel, and hence has the capability to depolarise the cell.

There seem to be three Na+ permeable channels in human taste cells:

i) one of which is selective/specific for Na+
ii) a fairly non-selective cation channel
iii) the voltage-gated Na+ channel that is crucial for action potential firing.

The fact that the channels are quite different can be demonstrated experimentally using selective
blocking agents.

 The Na+ specific channel can be blocked by the substance amiloride, whereas the non-
specific cation channel and the voltage-gated Na+ channels cannot.
 The non-specific cation channel can be blocked by the substance cetylpyridinium chloride
(CPC), whereas this substance does not affect the other two types of Na+ channel.
 The voltage-gated Na+ channel can be blocked by local anaesthetic (L.A) solutions, while
L.A. does not affect the other channels.

When a salty solution is applied to the tongue, the Na+ ions from it enter the Type III cells
through the first two two types of Na+ channels just described faster than the Na+ pump (Na+/K+
ATPase) can extrude them, and consequently there is an increase in intracellular Na+ and the cell
becomes depolarised (i.e., a receptor potential is generated). An action potential (or potentials)
follows, this is accompanied by an  Ca++ permeability, a Ca++ influx, and hence release of
transmitter (5-HT) at the synapses between the Type III cell and its associated afferent nerve
fibre.

Type I cells may also respond to salty stimuli.

Acid
The natural expectation is that the effective stimulus for acid sensations is H+ ions, protons, in the
fluid bathing the tongue. It has been realised for quite some time, however, that things are not
quite as easy as that. For example, the sourness of an acid solution is not a simple function of the
proton concentration (pH) of the taste solution. The intensity of sourness of different organic and
inorganic acids is poorly correlated with their pH: for example, acetic acid at pH 3.9 evokes a
strong acid taste in humans, but HCl at this pH hardly tastes acid at all. The implication of this is
that other components of the tastant molecule (the acid) contribute to the transduction process.
Recent research has identified two proteins on Type III cells that work together to form a
functional H+ channel. They are the polycystic-kidney disease-2-like 1 ion channel and
polycystic-kidney disease-1-like 3 ion channel (PKD2L1/ PKD1L3). These two proteins belong
to a family of proteins that act as transient receptor potential (TRP) channels. Thus influx of H+
ions into the cell down their concentration gradient via this channel brings about a short-lived
depolarising current – in other words, a receptor potential, and an action potential follows. This is
as a result of protons inactivating (closing) K+ channels thereby leading to cellular
depolarisation. This leads to opening of voltage-gated Ca++ channels, an influx of Ca++, and
transmitter (5-HT) release.

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Taste qualities detected by Type II cells
Type II cells have the capability to express a series of genes that code for receptor proteins. There
are two families of these proteins, the Taste receptor type 1 (T1R) and the Taste receptor type 2
(T2R) families. Only the members of the T1R family form dimers. (A dimer is a combination of
two components, or sub-units). These dimers in the apical membrane of the cells create taste
receptors and operate as G-protein coupled receptors.

In the T1R family there are 3 separate genes, and these code for the proteins, T1R1, T1R2 and
T1R3. The combinations of these that have functional significance are as shown in the table
below.
Receptor properties Taste modality
T1R1 + T1R3 Amino acid sensing receptor Umami
T1R2 + T1R3 Broad-based sweet sensing receptor Natural and artificial sugars

Unlike the T1R family, there is as yet, no evidence that the T2R family forms dimers.

Sweet
The range of substances that taste sweet is very diverse. There are the natural sugars; for
example, monosaccharides such as glucose and fructose, and disaccharides such as sucrose,
lactose and maltose, but then there are also artificial sweeteners such as saccharine, aspartame
and the cyclamates. Despite the wide differences in the structures of these molecules there seems
to be just one receptor that is capable of detecting these substances, and it is the T1R2 + T1R3
receptor. Binding of a sweet tasting substance to the receptor brings about a complex series of
events, and these are set out in the flow chart - Figure 2.

The experimental evidence indicates that the Type II cells do not have direct functional
synaptic connections with afferent nerve fibres, but rather they have to communicate with
afferent fibres through either the Type III cells (using 5-HT) or through longer-distance diffusion
of ATP.

Umami
The umami receptor in the Type II cell membrane is another type of dimer, this time made up of
T1R1 + T1R3. It is excited by amino acids. Glutamate and aspartate are particularly strong
excitants. When either of these binds to the receptor it causes activation of the same sequence of
events as shown in Figure 2.

Bitter
As with sweet, many different kinds of substance can evoke a bitter taste. Included in bitter
tasting compounds are not only those we easily recognise as bitter, such as caffeine and quinine,
but also some amino acids, fatty acids, phenols, amines, esters, and some salts. It seems that
bitter-sensing taste cells only express subsets of the T2R family, rendering each group of cells
sensitive to only some, and not all bitter compounds. When a T2R receptor is activated by a bitter
substance, the sequence of events is as shown in Figure 2.

.
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 Binding of sweet substance to T1R2 + T1R3 receptor or umami substance to T1R1 + T1R3
receptor or bitter substance to T2Rs

Activation of the G-protein complex coupled to the receptor

Activation of phospholipase C

Increase in cytosolic inositide triphosphate (IP3)

Release of Ca++ from intracellular store

+
Influx of Na into Type II cell and an action potential results

Release of ATP from the Type II cell

The ATP released from Type II acts as an excitatory transmitter substance on nearby afferent
nerve endings (i.e. in the same taste bud). This excites the gustatory afferent nerves.

AND
The released ATP acts as an excitatory transmitter substance, and it acts on neighbouring Type
III cells

The neighbouring Type III cell is depolarised and there is an increase in its membrane
permeability to Ca++ ions: consequently Ca++ move into the cell down their concentration
gradient

The Ca++ influx triggers release of transmitter (5-HT) from the Type III cell, and there is
excitation of the associated afferent nerve ending and inhibition (negative feedback) of Type II
cells

Figure 2: Flow chart showing the transduction pathways of taste in Type II and Type III
cells

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RESPONSES OF TASTE AFFERENTS
In the section above the point was made that individual Type III and Type II cells are sensitive to
just one modality of taste, but individual gustatory (taste) afferents will not necessarily be taste
specific.
A summary of these facts is as follows.
 Type II cells communicate to taste afferents through neighbouring Type III cells
 ATP released from Type II cells might also bring about activation of taste afferent
endings in the same taste bud.
 An individual taste afferent may branch within a taste bud to supply more than one Type
III cell.
 An individual taste afferent may branch to supply more than one taste bud in the lingual
epithelium.

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INFORMATION EXTRACTION
Taste afferent nerve fibres transmit information from taste buds to the brain, however how they
specify different taste qualities e.g. sweet or bitter remains unclear. Three theories have been
proposed:
1. Labelled line theory - Specific populations of receptor cells are uniquely tuned to the
individual qualities and each quality is independently transmitted by specific ‘labelled’ neurones.
2. Across fibre theory - Any given neurone could transmit information for more than a
single quality (i.e. they are non-specific). Patterns of activity distinguish between the different
taste qualities.
Consider two afferents with different response profiles as given below.

Sweet Salt Acid Bitter

Afferent 1  
Afferent 2  

If the activity of both afferents is monitored by the CNS, it could analyse events in the following
way.
 If only afferent 1 is firing, the stimulus must be bitter.
 If only afferent 2 is firing, the stimulus must be salt.
 If afferents 1 and 2 are both firing, the stimulus must be acid.

3. A temporal code in which timing pattern of action potentials denote a particular quality.

CNS TASTE PATHWAYS

The primary afferent pathways from the taste buds (VII, IX and X cranial nerves) are all routed
through the nucleus of the solitary tract (left and right sides) in the medulla oblongata. From
here there are connections to the following.
 To the ventral posteromedial (VPM) part of the thalamus (crossed pathway), and then to
the facial area of the somatosensory cortical area (postcentral gyrus). It is here that taste
sensations are most likely generated.
 To the superior and inferior salivatory nuclei on both sides for reflex saliva secretion.

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  To the limbic cortex, via the thalamus. In developmental terms the limbic cortex is the
oldest part of the cerebral cortex, and it is involved with instinctive behaviour and
emotions. In addition to receiving inputs from gustatory pathways there are also olfactory
inputs. The limbic system plays a role in the sense of pleasure or disgust that come when
eating food.
 To the facial nerve. These are gusto-facial reflexes where the facial nerve responds to
tongue stimulation by taste stimuli. This is an innate characteristic of humans as it exists
even in newborns.
 To the hypoglossal nerve evoking reflex tongue movements.

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NEUROTROPHISM AND TASTE BUDS

If one of the gustatory nerves (VII, IX, X) supplying the tongue is damaged the taste buds in the
area of the tongue supplied by that nerve may begin to degenerate and then, over a period of
days, may disappear. This can occur, for example, if the lingual nerve is damaged during the
surgical removal of a mandibular third molar tooth (recall that at the point where the lingual
nerve is close to the mandibular molars it contains the gustatory afferents from the chorda
tympani nerve, and these innervate taste buds on the anterior 2/3 of the tongue.) If the damaged
nerve is able to regenerate, taste buds reappear in the relevant field of innervation.
This is an example of neurotrophism. Neurotrophins are molecular signals/proteins that are
important for nerve cell survival (neurotrophism) and axonal growth

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Material produced by Stephen Lisney, Lucy Donaldson and Margaret Gatumu