The Chemical Senses:

Taste and Smell

Smell and taste are chemical sensations

Taste is the gustatory sensation
Smell is the olfactory sensation
Taste drives appetite and protects us from poisons

 Taste is mainly a function of the taste buds in the mouth,

but it is common experience that ones sense of smell
also contributes strongly to taste perception.

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:
2 sodium receptors
2 potassium receptors
1 chloride receptor,
1 adenosine receptor
1 inosine receptor
2 sweet receptors
2 bitter receptors,
1 glutamate receptor
1 hydrogen ion receptor.

 the 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

Taste buds
aggregations of 30-100 individual elongated
"neuroepithelial" cells
50-60 microns in height
30-70 microns in width
embedded in specializations of surrounding epithelium,
termed papillae
Types of papillae:
...Fungiform, foliate, circumvallate, filiform

 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.

Taste buds
below the taste bud, taste cells are joined by tight
junctional complexes
At the base of the taste bud, afferent taste nerve axons
invade the bud and ramify extensively, each fibre
typically synapsing with multiple receptor cells within
the taste bud
located throughout the oral cavity, in the pharynx, the
laryngeal epiglottis and at the entrance of the
oesophagus
number of taste buds declines with age

 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.

Salt taste

Na+ ions enter the receptor

cells via Na-channels

These are amiloridesensitive Na+ channel

entry of Na+ causes a

depolarization

Ca2+ enters through

voltage-sensitive Ca2+
channels

neurotransmitter release
occurs and results in
increased firing in the
primary afferent nerve

Sour taste
Sour is acidic
H+ ions block K+
channels
Blocking of these
channels causes a
depolarization
Ca2+ entry, transmitter
release and increased
firing in the primary
afferent nerve

Sweet taste

glucose binding to receptors

activates adenylyl cyclase,
thereby elevating cAMP
This causes a PKA-mediated
phosphorylation of K+
channels, inhibiting them
Depolarization occurs
Ca2+ enters the cell through
depolarization-activated
Ca2+ channels
transmitter is released
increasing firing in the
primary afferent nerve

Bitter taste
Bitter substances cause
the second messenger
(IP3) mediated release of
Ca2+ from internal stores
(external Ca2+ is not
required)
The elevated Ca2+
causes transmitter
release
this increases the firing of
the primary afferent nerve

Umami taste
Umami is the taste of certain amino acids (e.g.
glutamate, aspartate and related compounds)
It was first identified by Kikunae Ikeda at the Imperial
University of Tokyo in 1909

Umami taste
Recently it has been shown that the metabotropic
glutamate receptor (mGluR4) mediates umami taste
Binding to the receptor activates a G-protein
this may elevate intracellular Ca2+

 Monosodium glutamate, added to many foods to

enhance their taste (and the main ingredient of Soy
sauce), may stimulate the umami receptors
But, in addition, there are ionotropic glutamate receptors
(linked to ion channels), i.e. the NMDA-receptor, on the
tongue.
When activated by these umami compounds or soy
sauce, non-selective cation channels open, thereby
depolarizing the cell.
Calcium enters, causing transmitter release and
increased firing in the primary afferent nerve

Modifying taste
Taste exhibits almost complete adaptation to a stimulus
perception of a substance fades to almost nothing in seconds

Taste can be suppressed by local anaesthetics applied to

the tongue
Amiloride, a blocker of epithelial Na channels, reduces
salt taste in humans
AMP may block the bitterness of several bitter tasting
agents

Taste map
the classical "taste map" is an over simplification
Sensitivity to all tastes is distributed across the whole
tongue and to other regions of the mouth where there
are taste buds (epiglottis, soft palate)
some areas are indeed more responsive to certain tastes
than others

Innervation
Taste receptor cells do not have an axon
Information is relayed onto terminals of sensory fibres by
neurotransmitter
These fibers arise from the ganglion cells of the cranial
nerves
Vll (facial - a branch called the chorda tympani)
lX (glossopharyngeal)

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 potentialthat 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.

Central pathways
Primary gustatory fibers synapse centrally in the medulla
(in a thin line of cells called the nucleus of the solitary
tract)
From there the information is relayed
1 to the somatosensory cortex for the conscious
perception of taste
2 to the hypothalamus, amygdala and insula, giving
the so-called "affective" component of taste
This is responsible for the behavioural response,
e.g. aversion, gastric secretion, feeding behaviour.

 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

through the chorda tympani

into the facial nerve
into the tractus solitarius in the brain stem.

Flavor
Flavor is a combination of
Taste
Smell
texture (touch sensation)
and other physical features
(eg. temperature)

Physiology of Smell

 odor molecules must be small enough to be volatile

(less than 300-400 relative molecular mass) so that
they can vapourise, reach the nose and then dissolve
in the mucus

 we can distinguish around 10,000 different smells

why we smell and the impact of smell on our everyday
life are poorly understood
anosmic (someone who has lost some or all of their
sense of smell)
some anosmics suffer from depression and their
quality of life is severely affected

Olfactory epithelium
Contains
sensory cells
Bowman's glands producing
the secretion that bathes the
surface of the receptors
This is an aqueous
secretion containing
mucopolysaccharides,
immunoglobulins, proteins
(e.g. lysozyme) and various
enzymes (e.g. peptidases)
pigmented-type of epithelial cell

Odorant Binding Proteins

facilitate the transfer of lipophilic ligands (odorants)
across the mucus layer to the receptors
increase the concentration of the odorants in the layer
relative to air
as a transporter, in which they would bind to a receptor
with the ligand and accompany it across the membrane
as a terminator, causing "used" odorants to be taken
away for degradation, allowing another molecule to
interact with the receptor
as a protector for the receptor, preventing excessive
amounts of odorant from reaching the receptor.

Mechanism of Excitation of the Olfactory Cells

The portion of each olfactory cell that responds to the
olfactory chemical stimuli is the olfactory cilia.
The odorant substance, on coming in contact with the
olfactory membrane surface, first diffuses into the
mucus that covers the cilia.
Then it binds with receptor proteins in the membrane of
each cilium.
Each receptor protein is actually a long molecule that
threads its way through the membrane about seven
times, folding inward and outward.
The odorant binds with the portion of the receptor
protein that folds to the outside.

 The insideof the folding protein is coupled to a so called

G-protein, itself a combination of three subunits.
On excitation of the receptor protein
an alpha subunit breaks away from the G-protein

immediately activates adenylyl cyclase, which is

attached to the inside of the ciliary membrane near the
receptor cell body.

The activated cyclase

converts many molecules of intracellular adenosine triphosphate into
cyclic adenosine monophosphate (cAMP)
cAMP activates another nearby membrane protein, a gated sodium
ion channel, that opens its gate
allows large numbers of sodium ions to pour through the membrane
into the receptor cell cytoplasm.
The sodium ions increase the electrical potential in the positive
direction inside the cell membrane,
exciting the olfactory neuron and transmitting action potentials into the
central nervous system by way of the olfactory nerve.

Odorant receptor neurons

bipolar neurons in the nasal epithelium
they are capable of regenerating
Contain cilia which project into the mucus (these contain
the receptor proteins)
axons that project to the olfactory bulb 10-100 axons
form up into bundles that penetrate the ethmoidal
cribriform plate and terminate in the olfactory bulb

Mitral cells
the principal neurons in
the olfactory bulb
There are about 50,000
of these cells in each
bulb
They have a primary
apical dendrite which
extends into a spherical
bundle of neuropil
called a glomerulus

 glomerulus receives the

input from the olfactory
receptor neurons
Their axons merge
together to form the
lateral olfactory tract
They possess collaterals,
involved in negative
feedback and positive
feed-forward

Other cells
Periglomerular cells
- are involved in lateral inhibition at the level of the
glomeruli
Granule cells
-inhibitory interneurones
Olfactory ensheathing cells
like glial cells

Central connections
Neurons from the lateral olfactory tract project to
Areas of the limbic system (amygdala, septal nuclei,
entorhinal cortex and hippocampus)
The septal nuclei and amygdala contain regions
known as the "pleasure centres
The hippocampus is concerned with motivational
memory (the association of certain stimuli with
food)

Projections are also sent to

the thalamus and to the
frontal cortex for recognition
There are many forward and
backward connections
between each of these brain
centres.

Neurotransmitters

Glutamate
Noradrenalin
Dopamine
GABA - inhibitory

 Transmission of Olfactory Signals into the Olfactory Bulb

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