CHEMICAL SENSES:

TASTE AND SMELL

* EMMANUEL A. GUTIERREZ, MD
 OBJECTIVES
▪ Discuss the functions of the different parts of the
olfactory system
▪ Enumerate characteristics of olfactory stimuli, and
know the current estimate of odorant molecules to
which the human olfactory system is sensitive to
▪ Explain how potentials are produces in the olfactory
cells and how the signals are transmitted to the
central nervous system
▪ Enumerate 2 general disorders of sense of smell, and
general mechanisms
 OBJECTIVES
▪ Discuss the functions of the different parts of the
gustatory system
▪ Enumerate the different primary sensations of tastes
▪ Explain how potentials are produced in the taste
receptor cells and how the signals are transmitted to
the CNS
▪ Enumerate 2 general disorders of sense of taste
INTRODUCTION: SMELL & TASTE

▪ Generally classified as visceral senses because of their

close association with gastrointestinal function
▪ Receptors are chemoreceptors that are stimulated by
molecules in solution in mucus in the nose and saliva in
the mouth
▪ Receptors are classified as exteroceptors since stimuli
arrive from an external source
▪ Strongly tied to primitive emotional and behavioral
functions of our nervous systems
SENSE OF SMELL
ANATOMIC FEATURE: SENSE OF SMELL
 OLFACTORY EPITHELIUM

▪ Covers an area of 5 cm2 in the roof of the nasal cavity

near the septum
▪ Contains 10 to 20 million bipolar olfactory sensory
neurons interspersed with glia-like supporting
(sustentacular) cells and basal stem cells.
▪ Said to be the place in the body where the nervous
system is closest to the external world (dorsal posterior
recess of the nasal cavity)
▪ New olfactory sensory neurons are generated by basal
stem cells as needed to replace those damaged by
exposure to the environment.
 OLFACTORY EPITHELIUM

▪ Each neuron has a short, thick dendrite that projects

into the nasal cavity where it terminates in a knob
containing 10 to 20 immotile cilia .
▪ The cilia are unmyelinated processes about 2 um long
and 0.1 um in diameter and contain specific receptors
for odorants (odorant receptors).
▪ The axons of the olfactory sensory neurons pass
through the cribriform plate of the ethmoid bone and
enter the olfactory bulbs
OLFACTORY BULBS
OLFACTORY BULBS
 OLFACTORY BULBS

* The axons of the olfactory sensory neurons (first

cranial nerve) contact the primary dendrites of the
mitral cells and tufted cells to form anatomically
discrete synaptic units called olfactory glomeruli.
* Contain periglomerular cells, which are inhibitory
neurons connecting one glomerulus to another
* Contain granule cells, which have no axons and make
reciprocal synapses with the lateral dendrites of the
mitral and tufted cells
 OLFACTORY CORTEX

* The axons of the mitral and tufted cells pass

*
 OLFACTORY CORTEX
1. Anterior olfactory nucleus – an integrative center connecting the
two olfactory bulbs through the anterior commissure; integration of
olfactory inputs; information-sharing
2. Piriform cortex - main area involved in olfactory discrimination;
electrical stimulation in humans produces olfactory sensations.
3. Olfactory tubercle – involved in various functions of limbic system;
also receives ascending dopaminergic inputs from the midbrains;
malfunction in this area leads to kinds of schizophrenia.
4. Amygdala – its corticomedial parts receive inputs from the main and
accessory olfactory bulbs; emotional responses to olf. stimuli.
5. Part of enthorhinal cortex – receives olfactory input and projects to
hippocampus; involved in olfactory memories.
 CHARACTERISTICS OF THE
OLFACTORY MOLECULE
1. Volatile – transported by air
2. Slighty water-soluble – penetrates hydrated mucous
layer of the nasal epithelium to reach the olfactory
cilia
3. Slightly lipid-soluble – to penetrate phospholipid
bilayer of olfactory receptor neurons to activate
those cells
4. Odorant molecules are generally small, from 3-4, to
18-20 carbon atoms; different structural
configurations give different odors.
PRIMARY SENSATIONS OF SMELL
Odor Class Example of Known Smells like
compounds
Floral Geraniol Roses

Ethereal Benzyl acetate Pears

Musky Muscone Musk

Camphor Cineole, camphor Eucalyptus

Putrid Hyrdogen sulfide Rotten eggs

Pungent Formic acid, acetic acid Vinegar

Pepperminty Peppermint
OLFACTORY THRESHOLDS
OLFACTORY DISCRIMINATION

▪ Humans can recognize more than 10,000 different odors

▪ However, determination of differences in the intensity of any
given odor is poor (concentration of an odor-producing
substance must be changed by about 30% before a difference
can be detected)
▪ Adapts very rapidly with continued exposure to an odorant
▪ The direction from which a smell comes may be indicated by
the slight difference in the time of arrival of odoriferous
molecules in the two nostrils
OLFACTORY DISCRIMINATION

How can humans discriminate 10,000 different odors?

1. Olfactory neurons utilize a large number of G
protein-coupled receptor molecules to detect a wide
range of odors.
2. The olfactory system recognizes and interprets
patterns of responses by neurons in the ascending
pathways
ABNORMALITIES IN ODOR DETECTION

* Anosmia – inability to smell

* Hyposmia or hypesthesia – diminished olfactory
sensitivity
* Hyperosmia – enhanced olfactory sensitivity
* Dysosmia – distorted sense of smell
SIGNAL TRANSDUCTION
SIGNAL TRANSDUCTION
MEMBRANE POTENTIAL AND ACTION
POTENTIAL IN OLFACTORY CELLS

* Membrane potential inside unstimulated olfactory cells

averages about –55 millivolts → generate continuous
action potentials at a very slow rate (once every 20
seconds up to two or three per second)
* Odorants cause depolarization, decreasing the
negative potential in the cell to −30 millivolts or less →
number of action potentials increases to 20 to 30 per
second
 VOMERONASAL ORGAN

▪ Found in In rodents and various other mammals; not well

developed in humans
▪ Patch of olfactory epithelium located along the nasal septum
(anterior third of the nasal septum in humans)
▪ Concerned with the perception of odors that act as
pheromones
▪ Project to the accessory olfactory bulb and from there
primarily to areas in the amygdala and hypothalamus that are
concerned with reproduction and ingestive behavior
 SNIFFING

▪ Amount of air reaching the olfactory epithelium is

greatly increased by sniffing
▪ An action that includes contraction of the lower part
of the nares on the septum, deflecting the airstream
upward.
▪ A semireflex response that usually occurs when a new
odor attracts attention.
 PAIN FIBERS IN THE NOSE

▪ Naked endings of many trigeminal pain fibers

▪ Stimulated by irritating substances and leads to the
characteristic "odor" of such substances as
peppermint, menthol, and chlorine
▪ Initiates sneezing, lacrimation, respiratory inhibition,
and other reflexes
ADAPTATION/DESENSITIZATION

▪ Continuously exposure to a even the most

disagreeable odor, perception of the odor decreases
and eventually ceases
▪ Mediated by Ca2+ acting via calmodulin on cyclic
nucleotide-gated (CNG) ion channels
▪ Special inhibitory cells in the olfactory bulb, the
granule cells
SENSE OF TASTE
TASTE BUDS
 TASTE BUDS

▪ Located in the mucosa of:

Epiglottis Pharynx
Palate Papillae of the tongue
▪ Fungiform papillae – flat anterior surface of the tongue;
moderate numbers of taste buds located at the top of the
papilla
▪ Circumvallate papillae – V shape on the posterior part of the
tongue; large number of taste buds along the sides of the
papilla
▪ Foliate papillae – Lateral surface of the tongue; moderate
numbers of taste buds along the sides of the papilla
 TASTE BUDS

▪ The specialized sense organ for taste (gustation) consists of

approximately 10,000 taste buds, which are ovoid bodies
measuring 50–70 um
▪ Each taste bud is composed of about 50 modified epithelial
cells:
➢ Taste cells/Receptor cells
➢ Supporting/sustentacular cells
➢ Basal cells
▪ Sensory neurons that respond to taste stimuli or tastants
▪ Life span of a taste bud = 10 days
TASTE BUDS

▪ Apical ends of taste cells

have microvilli/ taste
hairs that project into the
taste pore
▪ Each taste bud is
innervated by about 50
nerve fibers; each nerve
fiber receives input from
an average of five taste
buds.
 TASTE BUDS

▪ Four morphologically distinct types of cells within each

taste bud:
✓ basal cells,
✓ dark cells (Type I) – 60%
✓ light cells (Type II) - 30%
✓ Intermediate cells (Type III) -7%
PRIMARY SENSATIONS OF TASTE

▪ Sour –acids (H+)

▪ Salty – ionized salts mainly Na+
▪ Sweet - mainly organic substances e.g. sugars,
alcohols, glycols, aldehydes etc.
▪ Bitter - Long chain organic substances having nitrogen
and alkaloids (quinine, nicotine, caffeine etc.)
▪ Umami -  Glutamate and monosodium glutamate
 PRIMARY SENSATIONS OF TASTE
(13 Possible Chemical Receptors)

▪ 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
Transduction Mechanism of Salty
Substances

▪ Na+ enters amiloride- sensitive

Na+ channels known as ENac
causing membrane
depolarization, giving rise to a
receptor potential. Further
depolarization (which opens up
Ca++ channels) triggers release of
transmitter by gustatory receptor
cell, exciting adjacent afferent
neuron.
Transduction Mechanism by Acids

▪ H+ bind and block apical

K+ sensitive channels.
▪ H+ also enters amiloride-
sensitive epithelial sodium
(ENaC) ion channels
Transduction Mechanism of Sweet
Substances
▪ Glucose binds to membrane receptors,
coupled with G-proteins, activate adenylyl
cyclase, and raise [cyclic AMP]
➢ Cyclic AMP, via protein kinase A (PKA) phosphorylates K+
channels, reducing K+ conductance. This results in membrane
depolarization(producing a receptor potential).

▪ Artificial sweeteners activate phospholipase

C (PLC), producing second messenger
inositol triphosphate (IP3 ) and
diacylglycerol (DAG). IP3 causes release of
Ca 2+ from intracellular stores, while DAG
activates protein kinase C (PKC).
➢ The release of intracellular Ca 2+ stores from endoplasmic
reticulum results in release of transmitter substances.
Transduction Mechanism of Bitter
Substances

▪ Bitter-tasting molecules, such as

denatonium activate phospholipase C
(PLC), producing second messenger
IP3. IP3 increases intracellular Ca 2+
levels, leading to release of synaptic
transmitter, and activation of
gustatory nerve fiber.
▪ Denatonium also activates gustducin
(Ggust) and transducin, causing a
decrease in cyclic AMP; cAMP block of
cation channels is removed, resulting
in cation influx.
Transduction Mechanism of Umami

▪ Activation of truncated
metabotropic glutamate
receptor, mGluR4, in the taste
buds
▪ Glutamate may also activate
ionotropic glutamate receptors
to depolarize umami receptors
 SUMMARY: GENERAL
TRANSDUCTION MECHANISMS

1. Taste stimuli interact with the apical membrane,

either by:(a) binding to specific membrane receptors
or by (b) modulating apically located ion channels.

➢ Sour and salty substances directly induce receptor

potentials by entry into ion channels, or blockade of K+
channels.
➢ Bitter-tasting and sweet substances act via G-
protein-linked membrane receptors, activating a
second-messenger cascade involving cyclic AMP and/or inositol
triphosphate (IP3).
 SUMMARY: GENERAL
TRANSDUCTION MECHANISMS

➢ Umami taste depends on activation of glutamate receptors, both

ligand-gated (ionotropic) receptors and
metabotropic receptors.

2. The interaction usually leads to changes in

membrane conductance, generation of receptor
potentials, triggering of action potentials, Ca2+
influx, and transmitter release.
TASTE THRESHOLDS
DISCRIMINATION OF TASTE STIMULI
QUALITY AND INTENSITY

Sensitivity differs in
different areas, but all
tastes can be perceived
at most areas of the
tongue
DISCRIMINATION OF TASTE STIMULI
QUALITY AND INTENSITY

▪ Each nerve fiber in the gustatory nerves responds to more

than one taste stimuli.
▪ However, each fiber responds best to one of the four
primary taste qualities.
▪ The coding of a gustatory sensation (taste quality) is not a
simple, labeled-lined mechanism. It is a pattern of activities
in nerve fibers activated by a particular stimulus.
▪ Taste stimulus intensity discrimination is crude, as it takes a
30% increase in concentration before a difference in taste
stimuli intensity is detected. Note similarity with smell
discrimination
TASTE PATHWAYS
 ADAPTATION OF TASTE

Adaptation of the gustatory system (almost complete,

within a minute or so of continuous stimulation) results
from:

▪ Sensory adaptation (ADAPTATION at the level of the

taste bud cells)
▪ Central adaptation (adaptation at the level of higher-
order neurons in the CNS) – mechanism not known
 TASTE PREFERENCE AND CONTROL
OF THE DIET
▪ The phenomenon of taste preference located in the central
nervous system and not from a mechanism in the taste
receptors themselves .
▪ The following experiments demonstrate this ability of animals
to choose food in accord with the needs of their bodies:
➢ Adrenalectomized, salt-depleted animals = select drinking water
with a high concentration of sodium chloride in preference to
pure water
➢ Animal given injections of excessive amounts of insulin =
automatically chooses the sweetest food
➢ Parathyroidectomized animals automatically choose drinking
water with a high concentration of calcium chloride
 TASTE PREFERENCE AND CONTROL
OF THE DIET
▪ An important reason for believing that taste preference is
mainly a central nervous system phenomenon is that previous
experience with unpleasant or pleasant tastes plays a major
role in determining one’s taste preferences
▪ For instance, if a person becomes sick soon after eating a
particular type of food, the person generally develops a
negative taste preference, or taste aversion.
 FACTORS INFLUENCING TASTE
SENSATION

▪ Area of stimulation – threshold for taste decreases as

area of application of stimulus increases
▪ Temperature of substance – maximum sensitivity to
taste is obtained at 30-40 degree C
▪ Olfaction – as it affects flavor
▪ Individual variation – decrease in taste sensitivity in
older people due to atrophy of taste buds
 FACTORS INFLUENCING TASTE
SENSATION

▪ Sex – women are more sensitive to sweet and salt and

less sensitive to sour than men
▪ Adaptation – taste buds adapt quickly to a particular
taste
▪ Interaction between taste producing substances
affect taste sensation
For e.g. very sour lemon juice tastes good when mixed with sweet
sugar
AFFECTIVE NATURE OF TASTE

▪ Pleasantness and unpleasantness are called affective

attributes of a sensation. These make an individual
select or reject food.
▪ For e.g., sweet is unpleasant at very low conc but very
pleasant at high conc - sour and bitter are pleasant at
low conc but very unpleasant at high conc.
DISORDERS OF TASTE SENSATION

* Ageusia- absence of taste sensation- congenital or due to

lesion of facial or glossopharyngeal nerve.
* Hypogeusia- diminished taste sensation due to some diseases.
* Dysgeusia/Parageusia- altered taste sensation (metallic, salty,
foul or rancid taste; It is usually associated with temporal lobe
syndrome; temporary problem
* Familial dysautonomia- a very rare condition where a very
high conc of glucose fails to produce sweet sensation.
* Selective taste blindness- inherited Mendelian recessive trait.
They have a very high increase in threshold to bitter taste but
all other taste sensations are normal to them.
 RECENT DISCOVERIES

▪ Saccharin is 600 times as sweet as sucrose

▪ Miraculin - taste modifying protein obtained from
miracle fruit in Africa. - After eating this fruit sour
substance taste sweet. For eg lemon tastes like
orange
 SOURCES

• Hall, J.E. (2016). Guyton and Hall Textbook of Medical

Physiology, 13th Ed.
• Barret, K.E. et al (2016). Ganong’s Review of Medical
Physiology
• BRS Physiology, 5th Ed.