How do we sense odors ?
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The olfactory epithelium contains the
olfactory sensory neurons
Olfactory
sensory
neurons
project to the
olfactory bulb
Mucus protects
the olfactory
cilia, which
contain the
olfactory
receptor
molecules
Fig. 6.13
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What's unusual about the
olfactory system?
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Olfactory receptors have seven helical
transmembrane domains (I-VII)
They bind
odorants deep
in their interior
(red areas)
Structural
differences (mainly in
the yellow regions)
cause different
receptors to bind
different odorants
Fig. 6.14
Striedter, © 2016 Oxford University 4
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The family of olfactory receptor genes
contains many pseudogenes
~10% of the olfactory receptor genes in ancestral primates were pseudogenes
and many more became non-functional in the lineage leading to humans
Fig. 6.15
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Using dogs to smell out
medical conditions
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The Olfactory Transduction
Cascade
continue at top right
Fig. 6.16
Striedter, © 2016 Oxford University 7
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The Olfactory Transduction
Cascade
• The binding of an odorant to an olfactory
receptor activates a G protein called Golf, which
then activates adenylate cyclase.
• Activated adenylate cyclase raises intracellular
levels of cAMP, which opens cAMP-gated Na/Ca
channels.
• Opening the Na /Ca channels depolarizes the
cell and elevates intracellular Ca++ levels.
• Note the similarities (and differences) to the
phototransduction cascade.
• G protein coupled receptors all share some
fundamental similarities, but work through
different G proteins and interact with different
enzymes. 8
The axons of sensory neurons expressing
the same olfactory receptor converge in the
olfactory bulb
The blue-stained neurons all express the P2 type of olfactory receptor
whole-mount coronal section Fig. 6.17
Striedter, © 2016 Oxford University
Press
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Chemotopy: changes in odorant structure shift
neural activity systematically across the
olfactory bulb
• Rats were injected with radioactive 2-DG and
allowed to smell one of several odorants (pentanal,
hexanal, etc.).
• Shortly thereafter, the rats were sacrificed and
processed to determine where in the olfactory bulb
the radioactive 2-DG had accumulated, indicating
regions of high neuronal activity.
• The pseudocolored images reveal 2-DG uptake in a
digitally flattened glomerular layer after exposure
to aldehydes and alcohols with varying numbers of
carbon atoms.
• The areas of highest 2-DG uptake move
systematically across the bulb as carbon number
increases, revealing a chemotopic map.
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Chemotopy: changes in odorant structure shift
neural activity systematically across the
olfactory bulb
Neural activity was assessed by the uptake of 2-deoxyglucose (2-DG)
The olfactory
bulb was
digitally
flattened
Fig. 6.18
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Axon-less granule cells generate lateral
inhibition through dendrodendritic synapses
They are excited by mitral cells and, in turn, inhibit those same
mitral cells, as well as adjacent ones
Fig. 6.19
Striedter, © 2016 Oxford University
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Axon-less granule cells generate lateral
inhibition through dendrodendritic synapses
• Mitral cells have axons that project out of the
olfactory bulb, but they also release glutamate
from their dendrites onto the dendrites of
granule cells.
• The granule cells, in turn, release GABA back
onto the dendrites of mitral cells.
• These reciprocal synapses provide feedback
inhibition to activated mitral cells. They also
provide a mechanism through which an active
mitral cell can inhibit less active neighbors, a
phenomenon called lateral inhibition.
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Lateral inhibition in the olfactory bulb
sharpens odor representations
The most active mitral cells inhibit their less active neighbors,
which respond most strongly to different odorants
Fig. 6.20
Striedter, © 2016 Oxford University 14
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Brain Exercise
What are some similarities between
olfaction and vision?
What are the key differences?
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How do we sense odors ?
• The sense of smell is based on the binding of odorants to olfactory
receptor molecules in the cilia of olfactory sensory neurons, which
are located in the olfactory epithelium.
• Olfactory sensory neurons regularly die despite being protected by
nasal mucus, but they can be replaced even in adulthood. The
axons of olfactory sensory neurons project to the olfactory bulb.
• Olfactory receptor molecules are G protein-coupled receptors
resembling opsins. They are extremely diverse, especially in non-
humans, with different receptor types binding different odorant
epitopes.
• Olfactory sensory neurons expressing the same type of olfactory
receptor are scattered across the olfactory epithelium, but their
axons converge onto just a few glomeruli in the olfactory bulb.
Adjacent glomeruli tend to be activated by similar odorants.
• Reciprocal dendrodendritic synapses between mitral cells and
granule cells mediate feedback inhibition.
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Electronic odor sensors and
their potential applications
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