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II.

General principles of sensory physiology


The somatosensory System

III.

Chemical Senses

I.

IV.
V.

Vision
Hearing and Equilibrium

Lect. univ. dr. Loredana - Cristina MEREU


Laboratory of Biophysics & Med. Physics, Faculty of Physics,
'Alexandru Ioan Cuza' University of Iasi

Chemical Senses
Mechanical and chemical receptors sense the
bodys condition.
Three sensory systems associated with the nose
and mouth olfaction, taste, and the trigeminal or
general chemosensory system are dedicated to
the detection of chemicals in the environment.
All three of these chemosensory systems rely on
receptors in the nasal cavity, mouth, or on the face
that interact with the relevant molecules and
generate receptor and action potentials, thus
transmitting information about chemical stimuli to
appropriate regions of the central nervous system.

Chemical Senses

1. The olfactory system detects airborne


molecules called odorants.
In humans, odors provide information about food,
self, other people, animals, plants, and many other
aspects of the environment.
Olfactory information
can
influence
feeding
behavior, social interactions
and, in many animals,
reproduction.

Chemical Senses

2. The taste (or gustatory)


system detects ingested, primarily
water-soluble
molecules
called
tastants. Tastants
provide
information about the quality, quantity,
and safety of ingested food.
3. The trigeminal chemosensory
system provides information about
irritating or noxious molecules that
come into contact with skin or
mucous membranes of the eyes,
nose, and mouth.

Chemical Senses

Some sensory cells, called chemoreceptors,


contain membrane proteins that can bind to particular
chemicals in the extracellular fluid.
In response to this chemical interaction, the
membrane of the sensory neuron becomes depolarized,
leading to the production of action potentials.
Chemoreceptors are used
in the senses of taste and smell
and are also important in
monitoring
the
chemical
composition of the blood and
cerebrospinal fluid.

Chemical Senses

0.1 mm

The most sensitive chemoreceptors are on


sensory hairs of the male silkworm which detect sex
pheromones.

Olfactory system - Smell


In terrestrial vertebrates, the sense of smell, or
olfaction, involves chemoreceptors located in the
upper portion of the nasal passages. Humans detect
smells by means of olfactory neurons located in the
lining of the nasal passages. The axons of these
neurons transmit impulses directly to the brain via the
olfactory nerve.
Each
olfactory
receptor
is
specialized
for 1 odorant
molecule.

Olfactory system - Smell

Specialized neurons present in the olfactory


epithelium in the nose project cilia into a mucus layer.
The cilia are able to bind to odorant molecules the
binding triggers an AP which is transmitted to the
olfactory area of the olfactory bulb olfactory cortex.

Olfactory system - Smell

A terrestrial vertebrate uses its sense of smell in


much the same way that a fish uses its sense of taste to sample the chemical environment around it.
Because terrestrial vertebrates are surrounded by
air rather than water, their sense of smell has become
specialized to detect airborne particles (but these
particles must first dissolve in extracellular fluid before
they can activate the olfactory receptors).
The sense of smell can be extremely acute in
many mammals, so much so that a single odorant
molecule may be all that is needed to excite a given
receptor.

Olfactory system - Smell

From an evolutionary perspective, the


chemical senses - particularly olfaction - are
deemed the oldest sensory systems;
nevertheless, they remain in many ways the
least understood of the sensory modalities.
Although humans can detect only four(five)
modalities of taste, they can discern thousands of
different smells. New research suggests that there
may be as many as a thousand different genes coding
for different receptor proteins for smell.
The particular set of olfactory neurons that
respond to a given odor might serve as a fingerprint
the brain can use to identify the odor.

The Organization of the Olfactory System

The olfactory system is the most thoroughly


studied component of the chemosensory triad and
processes
information
about
the
identity,
concentration, and quality of a wide range of chemical
stimuli that we associate with our sense of smell.
These stimuli, called odorants, interact with
olfactory receptor neurons found in an epithelial sheet the olfactory epithelium - that lines the interior of the
nose.

The Organization of the Olfactory System

The axons arising from the receptor cells project


directly to neurons in the olfactory bulb, which in turn
sends projections to the pyriform cortex in the temporal
lobe as well as other structures in the forebrain.
The pyriform cortex
is
three
layered
archicortex - considered
to be philogenetically
older than the neocortex and thus represents a
specialized
processing
center
dedicated
to
olfaction.

The Organization of the Olfactory System

Projections from the pyriform cortex relay


olfactory information via the thalamus to association
areas of the neocortex. The olfactory tract also projects
to a number of other targets in the forebrain, including
the hypothalamus and amygdala. The further processing
that occurs in these various regions identifies the
odorant and initiates appropriate motor, visceral, and
emotional reactions to olfactory stimuli.
fMRI images showing
focal activity in the
regions of the olfactory
bulb, pyriform cortex, and
amygdala in a normal
human being passively
smelling odors.

Olfactory Perception in Humans

In humans, olfaction is often


considered the least acute of the
senses, and a number of animals
are obviously superior to humans
in their olfactory abilities.

Olfactory Perception in Humans

This difference may reflect the larger number of


olfactory receptor neurons (and odorant receptor
molecules) in the olfactory epithelium in many species
and the proportionally larger area of the forebrain
devoted to olfaction.

In a 70-kg human, the surface area of the olfactory


epithelium is approximately 10 cm2. In contrast, a 3-kg
cat has about 20 cm2 of olfactory epithelium.

Olfactory Perception in Humans

Since
the
number
of
odorants is very large, there have
been several attempts to classify
them in groups.
The most widely used classification was
developed in the 1950s by John Amoore, who divided
odors into categories based on:
- their perceived quality,
- molecular structure,
- and the fact that some people,
called anosmics, have difficulty
smelling one or another group.

Olfactory Perception in Humans

Amoore classified odorants as:

Pungent
Floral
Musky
Earthy
Ethereal
Camphor
Peppermint
Ether
Putrid

Olfactory Perception in Humans

These categories are still used to describe odors,


to study the cellular mechanisms of olfactory
transduction, and to discuss the central representation
of olfactory information. Nevertheless, this classification
remains entirely empirical.
Chemical
structure
and human perceptual
threshold
for
12
common odorants.
Molecules perceived at
low concentrations are
more
lipid-soluble,
whereas those with
higher thresholds are
more water-soluble.

Olfactory Perception in Humans

A further complication in rationalizing the


perception of odors is that their quality may change
with concentration.
For example, at low
concentrations
indole
has
a
floral
odor,
whereas
at
higher
concentrations it smells
putrid.
Although most people are able to
consistently identify a broad range of test
odorants, others fail to identify one or more
common smells.

Olfactory Perception in Humans

Such chemosensory deficits, called anosmias,


are often restricted to a single odorant, suggesting that a
specific element in the olfactory system, either an
olfactory receptor gene or genes that control expression
or function of a specific odorant receptor gene, is
inactivated.
Anosmias often target
perception
of
distinct,
noxious odorants. About 1
person in 1000 is insensitive
to butyl mercaptan, the foulsmelling odorant released
by skunks.

Olfactory Perception in Humans

More serious is the inability to detect hydrogen


cyanide (1 in 10 people), which can be lethal, or ethyl
mercaptan, the chemical added to natural gas to aid in
the detection of gas leaks.

A more radically diminished or distorted sense of


smell can accompany eating disorders, psychotic
disorders (especially schizophrenia), diabetes, taking
certain medications, and Alzheimers disease (all for
reasons that remain obscure).

Olfactory Perception in Humans

Perceptual thresholds in anosmic and normal


subjects for related organic chemicals. In anosmics,
these chemicals are only detected as irritants at
relatively high concentrations (parts per million, ppm); in
normal subjects, they are first detected at much lower
concentrations as odors. The numbers 18 stand for the
aliphatic alcohols from methanol to 1-octanol.
Perceptual
thresholds for three
additional common
irritants - phenylethyl
alcohol
(PEA),
pyridine (Pyr), and
menthol (Men).

Olfactory Perception in Humans

The ability to identify odors normally decreases


with age. If otherwise healthy subjects are challenged
to identify a large battery of common odorants, people
between 20 and 40 years of age can typically identify
about 5075% of the odors, whereas those between 50
and 70 correctly identify only about 3045%.
Normal decline in olfactory
sensitivity with age. The ability
to identify 80 common odorants
declines markedly between 20
and 70 years of age.

The Olfactory Epithelium and Olfactory Receptor Neurons

The transduction of olfactory information


occurs in the olfactory epithelium, the sheet of neurons
and supporting cells that lines approximately half of the
nasal cavities. (The remaining surface is lined by
respiratory epithelium, which lacks neurons and serves
primarily as a protective surface).

The Olfactory Epithelium and Olfactory Receptor Neurons

Diagram of the olfactory epithelium


showing the major cell types:
olfactory receptor neurons and their cilia,
sustentacular (supporting) cells (that detoxify
dangerous chemicals)
basal cells.

The Olfactory Epithelium and Olfactory Receptor Neurons

Basal cells regenerate new olfactory neurons to


replace dead or damaged cells. Olfactory neurons
typically live about one month.
Bowmans
glands
produce
mucus.
Nerve
bundles
of
unmyelinated
neurons and blood
vessels run in the
basal part of the
mucosa (called the
lamina propria).

The Olfactory Epithelium and Olfactory Receptor Neurons

The most important of the


olfactory epithelium cells is the
olfactory receptor neuron, a
bipolar cell that gives rise to a smalldiameter, unmyelinated axon at its
basal surface that transmits olfactory
information centrally.

The Olfactory Epithelium and Olfactory Receptor Neurons

These proteins have


seven
transmembrane
domains, plus a variable cell
surface
region
and
a
cytoplasmic
tail
that
interacts with G proteins.
G proteins, also known as
guanosine
nucleotidebinding proteins, are a
family of proteins involved
in transmitting signals from
a variety of different stimuli
outside a cell into the
inside of the cell.

The generic structure of putative olfactory odorant receptors.

The Olfactory Epithelium and Olfactory Receptor Neurons

Analysis of the complete


human genome sequence
has idenfied approximately
950 odorant receptor genes.
Each gene presumably
encodes an odorant receptor
that detects a particular set of
odorant molecules.
In rodents, genome analysis has
identified about 1500 different odorant
receptor genes. Thus, in mammals,
odorant receptors are the largest known
gene family, representing between 3 and
5% of all genes.

The Olfactory Epithelium and Olfactory Receptor Neurons

At its apical surface, the receptor neuron gives


rise to a single dendritic process that expands into a
knoblike protrusion from which several microvilli, called
olfactory cilia, extend into a thick layer of mucus.
The mucus that lines
the nasal cavity and controls
the ionic milieu of the
olfactory cilia is produced by
secretory
specializations
(called Bowmans glands)
distributed throughout the
epithelium. When the mucus
layer becomes thicker, as
during a cold, olfactory acuity
decreases significantly.

The Olfactory Epithelium and Olfactory Receptor Neurons

Two other cell classes,


sustentacular (supporting) cells,
the olfactory epithelium.
This entire apparatus - mucus
with neural and supporting cells mucosa.

basal cells and


are also present in
layer and epithelium
is called the nasal

The Olfactory Epithelium and Olfactory Receptor Neurons

(A) Scanning electron


micrograph of the human
olfactory
epithelium,
showing cell bodies of the
olfactory receptor neurons
(O) with their dendrites (D)
ending in cilia that form a
mat within the mucus layer
overlying the epithelium.
From the deeper aspect an
axon
(arrows)
arises,
forming bundles (Ax) in the
submucosa. Red blood
cells (r).

The Olfactory Epithelium and Olfactory Receptor Neurons

(B) High magnification


view of the distal dendritic
knob giving rise to
olfactory
cilia.
The
terminal web is visible
encircling
the
know
(arrows).

The Olfactory Epithelium and Olfactory Receptor Neurons

The superficial location of the nasal mucosa


allows the olfactory receptor neurons direct access to
odorant molecules. Another consequence, however, is
that these neurons are exceptionally exposed.
Airborne pollutants, allergens, microorganisms,
and other potentially harmful substances subject the
olfactory receptor neurons to more or less continual
damage.
Several
mechanisms
help maintain the integrity of
the olfactory epithelium in the
face of this trauma.

The Transduction of Olfactory Signals

Odorant transduction begins with odorant


binding to specific receptors on the external surface of
cilia. Binding may occur directly, or by way of proteins
in the mucus (called odorant binding proteins) that
sequester the odorant and are thought to shuttle it to
the receptor. Several additional steps then generate a
receptor potential by opening ion channels.

The Transduction of Olfactory Signals

Generation of receptor
potentials in response to odors
takes place in the cilia of receptor
neurons.

Thus, odorants evoke a


large inward (depolarizing) current
when applied to the cilia (left), but
only a small current when applied
to the cell body (right).

The Transduction of Olfactory Signals

The olfactory receptor neurons express an


olfactory-specific G-protein (Golf), which activates an
olfactory-specific adenylate cyclase. Both of these
proteins are restricted to the knob and cilia, consistent
with the idea that odor transduction occurs in these
portions of the olfactory receptor neuron. Stimulation of
odorant receptor molecules leads to an increase in
cyclic AMP (cAMP) which opens channels that permit
Na+ and Ca2+
entry
(mostly
thus
Ca2+),
depolarizing the
neuron.

The Transduction of Olfactory Signals

This depolarization, amplified by a Ca2+ activated Cl current, is conducted passively from the
cilia to the axon hillock region of the olfactory receptor
neuron, where action potentials are generated and
transmitted to the olfactory bulb.

The Transduction of Olfactory Signals

The receptor potential is reduced in magnitude


when
cAMP
is
broken
down
by
specific
phosphodiesterases to reduce its concentration.
At the same time, Ca2+ complexes with calmodulin
(Ca2+-CAM) and binds to the channel, reducing its
affinity for cAMP. Finally, Ca2+ is extruded through the
Ca2+/Na+ exchange pathway.

The Transduction of Olfactory Signals

Finally, like other sensory receptors, olfactory


neurons adapt in the continued presence of a stimulus.

The Transduction of Olfactory Signals

Adaptation is apparent subjectively as a


decreased ability to identify or discriminate odors during
prolonged exposure (e.g., decreased awareness of
being in a smoking room at a hotel as more time is
spent there).
Physiologically, olfactory receptor neurons
indicate adaptation by a reduced rate of action
potentials in response to the continued presence of an
odorant.

The Transduction of Olfactory Signals

Adaptation occurs because of:


(1) increased Ca2+ binding by calmodulin, which
decreases the sensitivity of the channel to cAMP;
and
(1) the extrusion of Ca2+ through the activation of
Na+/Ca2+ exchange proteins, which reduces the
depolarizing potential from Ca2+ activated Cl
channels.

The Transduction of Olfactory Signals

Transduction of stimulus energy into neural


activity by chemoreceptors and photoreceptors requires
intracellular second messengers.
A1. The olfactory cilia of the olfactory hair cell on
the mucosal surface bind specific odorant molecules
and depolarize the sensory nerve via a secondmessenger system. The firing rate signals the
concentration of odorant in the inspired air.

The Transduction of Olfactory Signals

A2. Chemoelectric transduction is produced when


the appropriate odorant binds to a receptor protein on
the cell membrane, which activates G proteins linked to
the receptor. Channel opening and depolarization in
olfactory receptors and certain gustatory receptors are
mediated by a second messenger (cAMP) stimulated by
G protein activation. A3. Receptor currents evoked by
the appropriate odorant.

The Transduction of Olfactory Signals

Olfactory sensory cell bioelectrogenesis


The transduction region in
grey - and signaling region in yellow.
Transduction occurs in the cilia, which
extend into the mucous layer.

A receptor-coupled second-messenger system


results in the opening of a cation-selective channel in the
ciliary membrane.

The Transduction of Olfactory Signals

Olfactory sensory cell bioelectrogenesis


The influx of cations depolarizes the cell membrane from
its resting level near 65 to 45 mV, in a graded
manner.
This depolarization spreads by passive current flow
through the dendrite to the soma.
A depolarization that reaches 45 mV is sufficient to
activate voltage-gated Na+ channels and initiate
impulse generation.
This Na+ current along with several varieties of voltagedependent K+ currents and a small Ca2+ current
produce one or more action potentials that are
propagated down the axon to the brain.

Olfactory Coding
How olfactory receptor
neurons
represent
the
identity and concentration
of a given odorant is a
complex issue that is
unlikely to be explained
solely by the properties of
the
primary
receptor
neurons.
Nevertheless, neurons
with specific receptors are
located in particular parts of
the olfactory epithelium.

Olfactory Coding

As in other sensory systems, the topographical


arrangement of receptor neurons expressing distinct
odorant receptor molecules is referred to as space
coding, although the meaning of this phrase in the
olfactory system is much less clear than in vision, where
a topographical map correlates with visual space.
The coding of olfactory information also has a
temporal dimension. Sniffing, for instance, is a periodic
event that elicits trains of action potentials and
synchronous activity in populations of neurons.
Information conveyed by timing is called temporal
coding and occurs in a variety of species.

Olfactory Coding

Responses of olfactory receptor neurons to


selected odorants. Neuron 1 responds similarly to
three different odorants. In contrast, neuron 2 responds
to only one of these odorants. Neuron 3 responds to
two of the three stimuli.
Downward
deflections represent
inward
currents
measured at a holding
potential of 55 mV.
In
vertebrates
responses to an odor
can be measured by an
electro-olfactogram.

Olfactory Coding

Responses of a single
olfactory receptor neuron to
changes in the concentration of
a single odorant, isoamyl
acetate.
The upper trace in each
panel (red) indicates the duration
of the odorant stimulus; the
lower
trace
the
neuronal
response. The frequency and
number in each panel of action
potentials increases as the
odorant concentration increases.

Olfactory Coding

Transducing and relaying odorant information


centrally from olfactory receptor neurons are only the
first steps in processing olfactory signals.
As
the
olfactory
receptor axons leave the
olfactory epithelium, they
coalesce (join) to form a large
number of bundles that
together
make
up
the
olfactory nerve (cranial
nerve I).

Olfactory Coding

Each olfactory nerve projects to the olfactory bulb


on that side, which lies on the ventral anterior aspect of
the ipsilateral forebrain.

The olfactory bulb provides the first stage of


synaptic processing of the sensory information in the
olfactory pathway.

Olfactory Coding

The olfactory bulb is comprised of several cell


layers:
Glomeruli
Mitral cells
Granule cells

Olfactory Coding

The most distinctive feature of the olfactory bulb is


an array of more or less spherical accumulations of
neuropil 100200 m in diameter called glomeruli,
which lie just beneath the
surface of the bulb and
receive
the
primary
olfactory axons.
Within
each
glomerulus, the axons of
the
receptor
neurons
contact the apical dendrites
of mitral cells, which are
the
principal
projection
neurons of the olfactory
bulb.

Olfactory Coding

Finally, granule cells,


which constitute the innermost
layer of the vertebrate olfactory
bulb, synapse primarily on the
basal dendrites of mitral cells
within the external plexiform
layer.
These cells lack an
identifiable axon, and instead
make
dendrodendritic
synapses on mitral cells.

Olfactory Coding

The olfactory system enables animals to detect


and discriminate between thousands of different odors.
The enormous number of olfactory receptors
allows for the ability to detect a vast array of odors.
Each olfactory receptor recognizes a subset of
chemical cues, with one odor activating more than one
receptor and one receptor recognizing more than one
odor.

Olfactory Coding

In the periphery, each olfactory neuron


expresses only one olfactory receptor (OR).
Neurons with the same OR
are distributed randomly in the
nasal epithelium but they all project
to the same glomerulus in the
olfactory bulb.
Thus, the activation of
different olfactory receptors leads to
the activation of different glomeruli
in the central nervous system.
Different combinations of
activated glomeruli represent
different smells.

Olfactory Coding

In mammals, one receptor is expressed per cell


and neurons with the same receptor project to the same
glomerulus, demonstrating that the logic of olfaction
has been maintained through evolutionary time.
In addition to the main olfactory pathway in
vertebrates, a parallel accessory pathway exists for
transmitting signals from less volatile odorous
compounds called pheromones (the accessory
olfactory system does not function in humans).
Humans do not detect pheromones and have
evolved other strategies to ensure appropriate detection
of mates and attackers.

Olfactory Coding

Higher order processing


begins in the olfactory bulb,
where
mitral/tufted
cells
synapse onto olfactory neurons
and transmit this information to
the olfactory cortex.
Periglomerular cells and
granule
cells
provide
inhibitory connections in the
bulb that shape olfactory
responses.

Olfactory Coding

Information is then relayed to five different brain


regions where it is ultimately translated into different
odor percepts and behavior.

Olfactory Coding

Diagram
summarizing
the
organization of the glomerular layer.

synaptic

Olfactory neurons with the same


receptor project to the same
glomerulus.
Mitral/tufted cells synapse onto a
single glomerulus.
Periglomerulur
cells
are
inhibitory
interneurons
that
synapse within and between
glomeruli.
Granule cells are inhibitory
interneurons
that
synapse
between mitral/tufted cells.

(a) The olfactory system begins in the peripheral structures of the nasal cavity.
(b) The olfactory receptor neurons are within the olfactory epithelium.
(c) Axons of the olfactory receptor neurons project through the cribriform plate
of the ethmoid bone and synapse with the neurons of the olfactory bulb.

Stem Cells and Olfactory Function


There is much current interest in stem cells and
the possibilities they raise for maintaining or repairing
brain function.
The olfactory system is unique in the adult brain in
being supplied by two sources of stem cells.
First is the olfactory epithelium, where new ORNs
arise from basal stem cells during development and
throughout the adult life of the animal. The second
example is the anterior migratory stream. This process
also occurs throughout adult life.
Further work is thus needed to gaining insights
into the fundamental problems of stem cell functions in
the brain.

Gustatory system - TASTE


The best definition of the gustatory system is that
it has specialized sensory cells in the periphery and
unique regions in the brain dedicated to sensory
processing.

The sensory cues detected by the gustatory


system are soluble chemicals, limiting detection to a
short range by direct contact with a chemical source.

The Organization of the Taste System

The taste system, acting in concert with the


olfactory and trigeminal systems, indicates whether
food should be ingested.
Once in the mouth, the chemical constituents of
food interact with receptors on taste cells located in
epithelial specializations called taste buds in the
tongue.

The Organization of the Taste System

Taste cells (the peripheral receptors) are found


in taste buds distributed on the dorsal surface of the
tongue, soft palate, pharynx, and the upper part of the
esophagus.

The Organization of the Taste System

The tongue is covered with small bumps, called


papillae, which contain taste buds that are sensitive to
chemicals in ingested food or drink. Different types of
papillae are found in different regions of the tongue.
The taste buds
contain specialized
gustatory receptor
cells that respond
to chemical stimuli
dissolved in the
saliva.

The Organization of the Taste System

The taste bud is a barrel-shaped structure


containing approximately 50100 taste cells.
Microvilli of the taste receptor cells project into an
opening in the epithelium, the taste pore, where they
make contact with gustatory stimuli.

The Organization of the Taste System

The taste cells transduce these stimuli and provide


additional information about the identity, concentration,
and pleasant or unpleasant quality of the substance.
This information also prepares the gastrointestinal
system to receive food by causing salivation and
swallowing (or gagging and regurgitation if the
substance is unpleasant).
Of course, food is not simply
eaten for nutritional value; taste
also depends on cultural and
psychological factors. How else can
one explain why so many people
enjoy consuming hot peppers or
bitter-tasting liquids such as beer?

The Organization of the Taste System

Information about the temperature and texture of


food is transduced and relayed from the mouth via
somatic sensory receptors from the trigeminal and
other sensory cranial nerves to the thalamus and
somatic sensory cortices.
Like the olfactory system, the taste system
includes both peripheral receptors and a number of
central pathways.

The Organization of the Taste System

Taste cells make synapses with primary sensory


axons that run in the chorda tympani and greater
superior petrosal branches of the facial nerve (cranial
nerve VII), the lingual branch of the glossopharyngeal
nerve (cranial nerve IX), and the superior laryngeal
branch of the vagus nerve (cranial nerve X),
whose cell bodies lie
within the cranial nerve
ganglia, to innervate the
taste buds in the tongue,
palate, epiglottis, and
esophagus, respectively.

The Organization of the Taste System

The central axons of these primary sensory neurons in


the respective cranial nerve ganglia project to rostral and
lateral regions of the nucleus of the solitary tract in the
medulla, which is also known as the gustatory nucleus of
the solitary tract complex.

Neural pathway for taste into the

gustatory cortex.

The Organization of the Taste System

Axons from the rostral (gustatory) part of the solitary


nucleus project to the ventral posterior complex of the
thalamus, where they terminate in the medial half of the
ventral posterior medial nucleus.
Neural pathway
Taste
impulses
travel
through nerves VII, IX and
X to a gustatory nucleus in
the medulla oblongata
(cross over) thalamus
gustatory cortex located in
the parietal lobe in the
mouth area.

Taste Perception in Humans

Most taste stimuli are nonvolatile, hydrophilic


molecules soluble in saliva.
Examples include salts such as NaCl needed for
electrolyte balance; essential amino acids such as
glutamate needed for protein synthesis; sugars such as
glucose needed for energy; and acids such as citric
acid that indicate the palatability of various foods
(oranges, in the case of citrate).
Bitter-tasting molecules, including plant alkaloids
like atropine, quinine, and strychnine, indicate foods
that may be poisonous. Placing bitter compounds in the
mouth usually deters ingestion unless one acquires a
taste for the substance, as for the quinine in tonic
water.

Taste Perception in Humans

Mammals are thought to perceive


only five taste modalities:
sweet
bitter
sour
salty
umami (from the Japanese word for delicious,
umami refers to savory tastes, including monosodium
glutamate and other amino acids)

Taste Perception in Humans

However, there are obvious limitations to this


classification. People experience a variety of taste
sensations in addition to these five, including astringent
(cranberries and tea), pungent (hot peppers and
ginger), fat, starchy, and various metallic tastes, to
name only a few.
In addition, mixtures of chemicals may elicit
entirely new taste sensations. But even though the
taste code defined by the five primary taste classes
is not yet fully understood, these tastes correspond to
distinct classes of receptors in subsets of taste cells.
Thus, taste perception is closely linked to the
molecular biology of taste transduction.

Taste Perception in Humans

Diagram of a tongue showing the distribution of


various taste bud populations, which are found in the
fungiform (F) papillae on the anterior tongue, the vallate
(V) and foliate (FO) papillae on the posterior tongue.
Our
complex
perception of taste is
the result of different
combinations
of
impulses in the sensory
neurons from these four
kinds of taste buds,
together with information
related to smell.

Taste Perception in Humans

The salty taste is produced by the effects of


sodium (Na+) and the sour taste by the effects of
hydrogen (H+).
Organic molecules that produce the sweet and
bitter tastes, such as sugars and quinine, respectively,
are varied in structure.

Taste Perception in Humans

Taste buds that respond best to specific tastes


are concentrated in specific regions of the tongue:
sweet at the tip
sour at the sides
bitter at the back
salty over most of the tongues surface.

Taste Perception in Humans

The taste system encodes information about the


quantity as well as the identity of stimuli. In general,
the higher the stimulus concentration, the greater the
perceived intensity of taste. Threshold concentrations
for most ingested tastants are quite high, however.
The effect of
smell on the sense of
taste can easily be
demonstrated
by
eating an onion with
the nose open and
then eating it with the
nose plugged.

Taste Perception in Humans

The concentration range for taste detection is


broad and depends on the nature of the chemical
stimulus.
At one extreme, taste cells detect sugars and
amino acids at very high concentrations (100
millimolar), allowing animals to detect only the most
caloric foodstuffs instead of food with little nutritional
value. At the other extreme, taste cells can also detect
minute amounts of noxious substances or toxins,
compounds that are harmful at very low concentrations.
Taste buds - collections of chemosensitive
epithelial cells associated with afferent neurons mediate the sense of taste in vertebrates.

Taste Perception in fish

In a fish, the taste buds are scattered over the


surface of the body. These are the most sensitive
vertebrate chemoreceptors known.
They are particularly sensitive to
aminoacids; a catfish, for example,
can distinguish between two different
amino acids at a concentration of less
than 100 parts per billion (1 g in
10,000 L of water)! The ability to taste
the surrounding water is very
important to bottom-feeding fish,
enabling them to sense the presence
of food in an often murky environment.

Taste Perception in flies

Like vertebrates, many arthropods also have taste


chemoreceptors. For example, flies, because of their
mode of searching for food, have taste receptors in
sensory hairs located on their feet. Each different
chemoreceptor detects a different type of food molecule.
The
sensory
hairs
contain
different
chemoreceptors that are able to detect sugars, salts,
and other molecules.
They can detect a
wide variety of tastes by the
integration of stimuli from
these chemoreceptors.

Transduction of Taste Signals

The major perceptual categories of taste - salty,


sour, sweet, umami, and bitter - are represented by five
distinct classes of taste receptors, which are found in
the apical microvilli of taste cells.
Salty and sour tastes are primarily elicited by
ionic stimuli such as the positively charged ions in salts
(like Na+ from NaCl), or the H+ in acids (acetic acid, for
example, which gives vinegar its sour taste). These ions
in salty and sour tastants initiate sensory transduction
via specific ion channels:
the amiloride-sensitive Na+ channel for salty taste,
and
an H+-sensitive, cation-selective channel for sour.

Transduction of Taste Signals

The receptor potential generated by the positive


inward current carried either by Na+ for salty or H+ for
sour depolarizes the taste cell. This initial
depolarization leads to the activation of voltage gated
Na+ channels in the basolateral aspect of the taste cell.
This additional depolarization activates voltagegated Ca2+ channels, leading to the release of
neurotransmitter from the basal aspect of the taste
cell and the activation of action potentials in ganglion
cell axons.

Molecular mechanisms of
taste transduction via ion
channels.

Transduction of Taste Signals

In humans and other mammals, sweet and amino


acid (umami) receptors are heteromeric G-proteincoupled receptors that share a common seven
transmembrane receptor subunit called T1R3, which is
paired with
the T1R2 seven-transmembrane receptor for
perception of sweet T1R2/T1R3
or
with the T1R1 receptor for amino acids T1R1/T1R3
The T1R2 and T1R1 receptors are expressed in
different subsets of taste cells, indicating that there are,
respectively, sweet- and amino acid-selective cells in the
taste buds.

Transduction of Taste Signals

Upon binding sugars or other sweet stimuli, the


T1R2/T1R3 receptor initiates a G-protein-mediated
signal transduction cascade that leads to the
activation of the phospholipase C isoform PLC2,
leading in turn to increased concentrations of inositol
triphosphate (IP3) and to the opening of TRP
channels (specifically the TRPM5 channel ~ Transient
receptor potential cation channel subfamily M member
5), which depolarizes the taste cell via increased
intracellular Ca2+ .
Transduction of amino acid stimuli via the
T1R1/T1R3 receptor also reflects G-protein-coupled
intracellular signaling leading to PLC2-mediated
activation of the TRPM5 channel and depolarization of
the taste cell.

Transduction of Taste Signals

For sweet tastants, heteromeric complexes of


the T1R2 and T1R3 receptors transduce stimuli via a
PLC2-mediated, IP3(inositol triphosphate)-dependent
mechanism that leads to activation of the TRPM5 Ca2+
channel.
For amino acids, heteromeric complexes of
T1R1 and T1R3 receptors transduce stimuli via the
same PLC2/IP3/TRPM5-dependent mechanism.

Molecular
mechanisms of taste
transduction via ion
channels
and
Gprotein-coupled
receptors.

Transduction of Taste Signals

Another family of G-protein-coupled receptors


known as the T2R receptors transduce bitter tastes.
Although the transduction of bitter stimuli relies on
a similar mechanism to that for sweet and amino acid
tastes, a taste cell-specific G-protein, gustducin, is
found primarily in T2R-expressing taste cells and
apparently contributes to the transduction of bitter tastes.
The remaining steps in bitter transduction are
similar to those for sweet and amino acids: PLC2mediated activation of TRPM5 channels depolarizes the
taste cell, resulting in the release of neurotransmitter at
the synapse between the taste cell and sensory ganglion
cell axon.

Transduction of Taste Signals

Bitter tastes are transduced via a distinct set of


G-protein-coupled receptors, the T2R receptor subtypes.
The details of T2R receptors are less well established;
however, they apparently associate with the taste cell
specific G-protein gustducin, which is not found in
sweet or amino acid receptor- expressing taste cells.
Nevertheless, stimulus-coupled depolarization for
bitter tastes relies upon the same PLC2/IP3/TRPM5dependent mechanism used for sweet and amino acid
taste transduction.

Molecular mechanisms of
taste transduction via ion
channels
and
G-proteincoupled receptors.

Basic components of sensory


transduction in taste cells.

Transduction of Taste Signals

Taste cells are polarized


epithelial cells with an apical and a
basolateral domain separated by
tight junctions.
Tastant-transducing channels
(salt and sour) and Gproteincoupled receptors (sweet, amino
acid, and bitter) are limited to the
apical domain.
Intracellular signaling components that are
coupled to taste receptor molecules (G-proteins and
various second messenger-related molecules) are also
enriched in the apical domain.

Transduction of Taste Signals


Basic components of sensory transduction in taste cells.

Voltage - regulated
Na+, K+, and Ca2+ channels
that mediate release of
neurotransmitter
from
presynaptic specializations
at the base of the cell onto
terminals
of
peripheral
sensory afferents are limited
to the basolateral domain, as
is endoplasmic reticulum that
also modulates intracellular
concentration
and
Ca2+
contributes to the release of
neurotransmitter.

Transduction of Taste Signals


Basic components of sensory transduction in taste cells.

The neurotransmitter
serotonin, among others, is
found in taste cells, and
serotonin
receptors
are
found on the sensory
afferents.
Finally, the TRPM5
channel, which facilitates Gprotein-coupled
receptormediated depolarization, is
expressed in taste cells. Its
localization to apical versus
basal domains is not yet
known.

Transduction of Taste Signals

A taste receptor cell responding to:


Na+ salt;

sweet solutes;

acid and sour solutes.

Transduction of Taste Signals

The binding of the receptor to a taste molecule


triggers the entry of calcium in the cell release of
neurotransmitter in a synapse with a neuron.

Transduction of Taste Signals

Neural Coding in the Taste System

In the taste system,


neural coding refers to
the way that the
identity,
concentration,
and
hedonic
(pleasurable
or
aversive)
value of tastants is
represented
in
the
pattern
of
action
potentials relayed to the
brain.

Neural Coding in the Taste System

Neurons in the taste system (or in any other


sensory system) might be specifically tuned to
respond with a maximal change in electrical activity to a
single taste stimulus.
Such tuning is thought to rely on specificity at the
level of the receptor cells, as well as on the
maintenance of separate channels for the relay of this
information from the periphery to the brain.
This sort of coding scheme is referred to as a
labeled line code, since responses in specific cells
presumably correspond to distinct stimuli.

Neural Coding in the Taste System

The segregated expression of sweet, amino acid,


and bitter receptors in different taste cells is consistent
with labeled line coding.

Specificity in peripheral
taste coding supports the
labeled line hypothesis.
(AC) Sweet (A), amino
acid (B), and bitter (C)
receptors are expressed
in different subsets of
taste cells.

Neural Coding in the Taste System

(DE) The gene for the TRPM5 channel can be


inactivated, or knocked out, in mice (TRPM5/) and
behavioral responses measured with a taste preference
test.

The mouse is presented with two drinking spouts, one


with water and the other with a tastant; behavioral
responses are measured as the frequency of licking of
the two spouts.

Neural Coding in the Taste System

For pleasant tastes like sweet (sucrose; D) or


umami (glutamate; E) control mice lick the spout with
the tastant more frequently, and higher concentrations
of tastant leads to increased response (blue lines).

In TRPM5/ mice, this behavioral response (i.e., a


preference for the tastant versus water) is eliminated at
all concentrations (red lines).

Neural Coding in the Taste System

(F) For an aversive tastant like bitter quinine,


control mice prefer water. This behavioral response
which is initially lowis further diminished with higher
quinine concentrations (blue line).

Neural Coding in the Taste System

Inactivation of TRPM5 also eliminates this


behavioral response, regardless of tastant concentration
(red line).
(GI) When the PLCb2 gene is knocked out,
behavioral response to (G) sucrose, (H) glutamate, and
(I) quinine are eliminated (red lines).

Neural Coding in the Taste System

When PLCb2 is re-expressed only in T2Rexpressing taste cells, behavioral responses to sucrose
and glutamate are not rescued (dotted green lines in G
and H); however, the behavioral response to quinine is
restored to normal levels (compare the blue and dotted
green lines in I).

Trigeminal Chemoreception
The

third of the
major
chemosensory
systems, the trigeminal
chemosensory
system,
consists of polymodal
nociceptive neurons and
their
axons
in
the
trigeminal nerve (cranial
nerve V) and, to a lesser
degree,
nociceptive
neurons whose axons run
in the glossopharyngeal
and vagus nerves (IX
and X).

Trigeminal Chemoreception
These neurons and their associated endings are
typically activated by chemicals classified as irritants,
including air pollutants (e.g., sulfur dioxide), ammonia
(smelling salts), ethanol (liquor), acetic acid (vinegar),
carbon dioxide (in soft drinks), menthol (in various
inhalants sensation), and capsaicin (the compound in
hot chili peppers that elicits the characteristic burning
sensation).

Trigeminal Chemoreception

Irritant-sensitive polymodal nociceptors alert the


organism to potentially harmful chemical stimuli that
have been ingested, respired, or come in contact with
the face, and are closely tied to the trigeminal pain
system.
Trigeminal
chemosensory
information from the face, scalp,
cornea, and mucous membranes of
the oral and nasal cavities is
relayed via the three major sensory
branches of the trigeminal nerve:
the ophthalmic, maxillary, and
mandibular.

Trigeminal Chemoreception

The central target


of these afferent axons
is the spinal component
of
the
trigeminal
nucleus, which relays
this information to the
ventral posterior medial
nucleus of the thalamus
and thence to the
somatic
sensory
cortex and other cortical
areas
that
process
facial irritation and
pain.

Internal Chemoreceptors
Sensory receptors within the body detect a
variety of chemical characteristics of the blood or fluids
derived from the blood, including cerebrospinal fluid.

Included among these receptors are the


peripheral chemoreceptors of the aortic and carotid
bodies and the central chemoreceptors in the
medulla oblongata of the brain.

Internal Chemoreceptors

Peripheral
chemoreceptors of
the
aortic
and
carotid bodies are
sensitive primarily to
plasma pH.
Central
chemoreceptors in
the
medulla
oblongata of the
brain are sensitive to
the
pH
of
cerebrospinal fluid.

Internal Chemoreceptors

When the breathing rate is too low, the


concentration of plasma CO2 increases, producing more
carbonic acid and causing a fall in the blood pH.
The carbon dioxide can also enter the
cerebrospinal fluid and cause a lowering of the pH,
thereby stimulating the central chemoreceptors. This
chemoreceptor stimulation indirectly affects the
respiratory control center of the brain stem, which
increases the breathing rate.
The aortic bodies can also respond to a lowering
of blood oxygen concentrations, but this effect is
normally not significant unless a person goes to a high
altitude.

Internal Chemoreceptors

Chemical Senses - Summary

The chemical senses - olfaction, taste, and the


trigeminal chemosensory system - all contribute to
sensing airborne or soluble molecules from a variety
of sources. Humans and other mammals rely on this
information for behaviors as diverse as attraction,
avoidance, reproduction, feeding, and avoiding
potentially dangerous circumstances.
Each of the approximately
10,000 odors that humans recognize
(and an undetermined number of
tastes and irritant molecules) is
evidently encoded by the activity of a
distinct population of receptor cells
in the nose, tongue, and oral cavity.

Chemical Senses Summary

Receptor neurons in the olfactory epithelium transduce


chemical stimuli into neuronal activity via the stimulation of Gprotein-linked receptors; this interaction leads to elevated levels
of second messengers such as cAMP, which in turn open cationselective channels. These events generate RP in the membrane
of the olfactory receptor neuron, and ultimately AP in the afferent
axons of these cells.
Taste receptor cells, in
contrast, use a variety of mechanisms
for transducing chemical stimuli.
These include ion channels that are
directly activated by salts and amino
acids, and G-protein-linked receptors
that activate second messengers.
For both smell and taste, the spatial and temporal patterns
of action potentials provide information about the identity and
intensity of chemical stimuli.

Chemical Senses Summary

Olfaction, taste, and trigeminal chemosensation all


are relayed via specific pathways in the CNS.
Receptor neurons in the olfactory system project
directly to the olfactory bulb. In the taste system,
information is relayed centrally by cranial sensory
ganglion cells to the solitary nucleus in the brainstem. In
the trigeminal chemosensory system, information is
relayed via trigeminal ganglion cell projections to the
spinal trigeminal nucleus in the brainstem.
Each of these structures project in turn to many
sites in the brain that process chemosensory
information in ways that give rise to some of the most
sublime pleasures that humans experience.

http://www.nature.com/nature/journal/v
486/n7403_supp/full/486S2a.html

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