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The Chemical Senses: Taste and Smell

Gustation: The sense of taste

Olfaction: The sense of smell
Odor: A general smell sensation of a particular quality
The cake has a chocolate odor
Odorant: Any specific aromatic chemical
You were given the odorant menthol to smell


We have an innate ability to detect bad smells
One day-old-babies will scrunch up their noses if exposed
to a rotten egg smell.
Identical twins have the same body odor:
Dogs can distinguish between the smell of T-shirts worn by
non-identical twins but not identical twins.
Moths can only smell a single moleculethe moth pheromone.
We can smell happiness:
Women could discriminate between armpit swabs taken
from people watching happy and sad films (Men were
less good at this).
Newborn babies will recognize their mothers odor.
Will prefer to nurse from Mom over other nursing mothers.

Not every chemical is an odorant.

tertiary butyl mercaptan

(a): Left smells like menthol and right smells like green bell peppers.
(b): Two molecules that dont have a smell: methane and carbon monoxide. Never evolved
sensitivity to these: not too much fossil fuel burning back in the day.
Tertiary-butyl mercaptan (smells like rotten eggs) added by the gas company to natural gas.
Not because these are too smallanisole and benzaldehyde are odorants (top, right).
Structure is kingR-carvone and S-carvone (mirror images) have completely different smells.

The human olfactory apparatus

Unlike other senses, smell is tacked onto an organ with another purposethe nose

Primary purpose: to filter, warm, and humidify air we breathe.

Nasal cavity contains small ridges and olfactory epithelium.
Olfactory epithelium: contains the cells that detect odorants in the inspired air.
Cribriform plate: A bony structure riddled with tiny holes, at the level of the eyebrows, that separates
the nose from the brain.
Axons from olfactory sensory neurons pass through the tiny holes to enter the brain.
Anosmia: The total inability to smell, most often resulting from head trauma
A hard blow to the front of the head can cause the cribriform plate to be jarred back or fractured,
slicing off the fragile olfactory neurons.
Neurons will regenerate after a few weeks.

Olfactory epithelium: The retina of the nose

Three types of cells:
Olfactory sensory neurons (OSNs): The main cell type in the olfactory
epithelium. These are the cells that actually detect odorants.
Supporting cells: provide metabolic and physical support for the
olfactory sensory neurons.
Basal cells: precursor cells to olfactory sensory neurons.

OSNs project to glomeruli.

Each glomerulus receives
input from one type of OSN.

Compared with other mammals, humans dont smell very well (good?)
The olfactory bulb
is proportionally
smaller in humans
than other
Humans have smaller olfactory bulbs and fewer receptors than rats and dogs
(despite our larger size).

Humans are good at detecting a number of compounds at

low concentrations.
It has been estimated that humans can smell about 10,000
different odors.
The smell of coconuts, bell peppers, urine and roses all arise
from the presence of a specific molecule that our olfactory
system can detect.

Seven common smells are used

to test for anosmia: the loss of (or
reduced) sense of smell:
baby powder, chocolate, coffee,
cinnamon, mothballs, peanut
butter and soap.
Anosmiacs are often restricted to
a single odorant suggesting a
specific element in the olfactory
system is inactivated, usually
because of a defective odorant
receptor gene.

Figure 15.4 Normal decline in olfactory sensitivity with age

The ability to identify odors

declines with age
Some diseases are associated with
decreased olfactory sensitivity.
associated with decreased
olfactory sensitivity---for
unknown reasons.

Olfactory neuron is
composed of several
The cell is bipolar in
nature with a
centrally located cell
body and an
axon extending in
one direction toward
the brain and a
dendrite that
projects toward
Bowmans gland makes mucus: odorants are
dissolved in the mucus and can then be detected
by olfactory cilia.

mucus layer that

ends with several
non-motile cilia.

Olfactory cilia are hair-like protrusions on the dendrites of OSNs.

Contain receptor sites for odorant molecules. These are the first
structures involved in olfactory signal transduction.
Application of the correct odorant to the cilia of an OSN leads to the
generation of an action potential that leads to the brain.
Cell only responds when odorant is applied
to cilia.

Olfactory sensory neuron (OSNs) vary in their responsiveness

to different odorants. Cells are tuned to different odorants and
have an odorant profile.
Odorant 1



Odorant 2

Odorant 3

Axons of OSNs make up cranial nerve I

Olfactory neurons synapse in glomerulus with mitral cells.

Each mitral cell receives input from one glomerulus.
Projects to amygdala-hippocampal complex.
Amygdala detects pleasant or unpleasantness
of smell.
Hippocampus mediates storage of memory
of smell.
Projection to olfactory cortex combines with other sensations
(taste, texture, vision) to give perception of flavor.

Olfactory Physiology
Limbic system: The encompassing group of neural
structures that includes the olfactory cortex, the
amygdala, the hippocampus, the piriform cortex, and
the entorhinal cortex.
Involved in many aspects of emotion and memory.
Olfaction is unique among the senses for its direct
and intimate connection to the limbic system.
Olfactions unique connection to the limbic system
explains why scents tend to have such strong emotion

Humans can track scents like dogs

We can be trained (just like a dog).

Dog tracks the scent
of a pheasant dragged
along the green line.

Human tracks the

scent of chocolate

Olfactory receptor cells are different from all other

sensory receptor cells: They are not protected from
the atmosphere by any barrier and make direct
contact with the brain.
Contrast with visual receptors that are protected by
cornea, hearing receptors protected by eardrum,
taste buds are buried in papillae.
Therefore, many drugs can be inhaled directly to
the brain through the cribriform plate.
OSN axons are among the thinnest and slowest in
the body (unmyelinated, but OK because brain is
so close).
Takes longer to perceive odors compared to
other senses

Leading theory of olfactory perception:

Shape-pattern theory: Different scentsas a function of odorant-shape
to OR-shape fitactivate different arrays of olfactory receptors in the
olfactory epithelium.
These various arrays produce specific firing patterns of neurons in
the olfactory bulb, which then determine the particular scent we
Suppose there were six types of odorants and six types of receptors: blue and orange
triangles, blue and orange squares and blue and orange saw-toothed.
Blue triangle can bind to either triangle-shaped receptor (blue or orange) but not to the
square or saw-toothed receptors.

Binding to blue receptor gives larger response (more exact fit).

Another example
of shape-pattern theory.
From Chemicals
to Smells
More complex structure (blue triangle bound to orange square).
It can bind to either triangle or square receptors of either color.
Size of the response will be a function of the best fit.
Blue-to-orange and orange-to-blue are not as good as
blue-to-blue or orange-to-orange. But more complex structure
gives a wider range of responses.

Odor mixtures
We rarely smell pure odorants. Rather, we smell mixtures.
How do we process the components in an odorant mixture?
Two possibilities:
Analysis: Example from auditory mixtures. Chord of three notes
can played together and we perceive them all, but we can
detect each individual note.
Synthesis: Example from color mixtures. Mixing red and green
lights results in yellow light, but we cannot separately perceive
the red and green in the yellow.

Odor imagery
Humans have little or no ability to conjure odor
We do not think in smell very well.
We do not imagine smells very welldreams
with olfactory sensations are very rare.

Method for threshold detection of odors.

Staircase method: method for determining
the concentration of a stimulus required for
detection at a threshold level.
Stimulus is presented in increasing
concentrations until detection is
Then, concentration is decreased until
detection ceases.
Ascending and descending sequence
is repeated several times and
concentrations at which reversals
occur are averaged to determine
threshold detection.

Method for discrimination of odors:

Triangle test: participant is given three odors to smell, two of which are the
same and one is different.
Participant must identify the odd odor.
The order of the three odors is varied and tested several times to
increase accuracy.

same, same, different

same different, same
different, same, same

Sense of smell is essentially a change detector.
Examples: Walking into a bakery and can only smell
fresh bread for a few minutes.

Three types of adaptation:

Receptor adaptation: The biochemical phenomenon that

occurs after continuous exposure to an odorant, whereby the
receptors stop responding to the odorant and detection

Cross-adaptation: The reduction in detection of an odorant

following exposure to another odorant.
Presumed to occur because the two odors are similar
enough that they share one or more olfactory receptors for
their transduction.

Do these five fragrances

smell the same?
No, but because of olfactory
cross-adaptation, if youve
smelled four of them in
succession, the subtleties of
the fifth one may be lost to
your nose.

Cognitive adaptation: The psychological process by

which, after long-term exposure to an odorant, one is
no longer able to detect that odorant or has very
diminished detection ability.
Someone who wears perfume every day
cannot smell it and might put a lot on.
If you worked in a bakery, you wouldnt smell
bread every time you went to work.

Odor Hedonics
The liking dimension of odor perception, typically measured with scales pertaining to
an odorants perceived familiarity and intensity.

Familiarity: we tend to like odors weve smelled many times before.

Intensity has a more complicated relationship with odor liking:
Inverted U-shape function for pleasant smells
Linearly decreasing function for initially tolerable but not necessarily
unpleasant smells.

Rose smell

Fishy smell

Taste: its innate

Infants behavior and facial expressions reveal inborn preferences for
certain foods.

Different flavored foods placed on tips of infants tongues:

Sweet food evokes a smile-like expression followed by sucking.
Sour produces pursing and protrusion of lips.
Bitter produces gaping, movements of spitting, and sometimes vomiting movements.

Retronasal olfactory sensation: The sensation of an odor that

is perceived when chewing and swallowing force an
odorant in the mouth up behind the palate into the nose.
Such odor sensations are perceived as originating from
the mouth, even though the actual contact of odorant
and receptor occurs at the olfactory epithelium.
Flavor: The combination of true taste (e.g. sweet, salty, sour)
and retronasal olfaction.

Four kinds of papillae on the tongue:

1. Filiform papillae: Small structures on the tongue that provide
most of the bumpy appearance. Have no taste function.
2. Fungiform papillae: Mushroom-shaped structures (maximum
diameter 1 mm) that are distributed most densely on the
edges of the tongue, especially the tip. An average of six
taste buds per papilla are buried in the surface.
3. Circumvallate papillae: Circular structures that form an
inverted V on the rear of the tongue (three to five on each
side). Mound-like structures surrounded by a trench. Much
larger than fungiform papillae.
4. Foliate papillae: Folds of tissue containing taste buds.
Located on the rear of the tongue lateral to the circumvallate
papillae, where the tongue attaches to the mouth.

Taste buds and taste receptor cellsnot on the surface of the papillae, but
on the sides.
Taste pore lets in saliva that has dissolved tastant molecules.
Taste cells (aka taste receptor cells) make up taste bud.
Microvilli: Slender projections on the tips of taste receptor cells that
extend into the taste pore and have sites that detect tastants.

Tastant: Any stimulus that can be tasted.

Tastants can be divided into two categories:
1. Small, charged particles that taste salty or sour.
Ion channels in microvilli membranes allow some types of charged
particles to enter but not others.

2. Larger molecules that taste sweet, bitter or savory (umami).

These are detected via G protein-coupled receptors (GPCRs).

The different receptor mechanisms for ionic stimuli and those

using a lock-and-key.
Five basic tastes:
Salty: Na+ ions
Sour: H+ ions (acidic)
Bitter: Quinine, coffee (often
poisonous substances)
Sweet: Sucrose, fructose, glucose
and also artificial sweeteners all
activate the same receptor
Umami: savory monosodium
glutamate (MSG)

Survival value of taste

Taste is a system for detecting nutrients and antinutrients
Bitter: Might signal poisons (threshold for
detection is very low: need to respond to low
levels of poisons)
Sour: Configured to detect acidic solutions that
might harm the body (spoiled food)
Sweet and Salty: Our bodies need sodium and
sugar to survive (threshold is relatively high:
need these for survival, dont want to be
satisfied with low levels)

Pleasure and retronasal versus orthonasal olfaction

Retronasal olfaction: olfaction through the mouth.
Orthonasal olfaction: olfaction through the nostrils.
Do we learn to like or dislike smells separately for
retronasal versus orthonasal olfaction? Possibly
Example: Many people like the smell of freshly
cut grass, but wouldnt want to eat it.
However, if an aversion is acquired retronasally, it
usually shows up orthonasally as well.
Example: Becoming sick from eating fish and
then disliking even the smell of fish.