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Olfaction – the primal sense, forgotten sense, smaller role for humans
Significance: alarm system (fire, smokers), flavor, communication (pheromones), social context (family,attractiveness)
Mercaptan – smelly chemical + natural gas = smelly gas leak
Pathway: Air passes olfactory epithelium with chemosensors neurons in olfactory bulb sensory cortex
OSNs (olfactory sensory neurons) – specialized cells that detect gaseous molecules and produce an electrical signal
Chemo-electrical signal transduction – turning chemical signal into electrical signal
Transduction all located within sensory cilia of OSN
Odorants
Volatile chemical compounds carried by inhaled air to olfactory epithelium
Humans can smell 4K-10K odors
Size of epithelium related to sense of smell = larger size means greater sense (humans 10 cm2 vs dogs 170 cm2)
Molecular properties: water solubility, volatile, vaporizable, low polarity / non-ionic, lipophilicity, surface activity
Each odorant activates a unique set of olfactory receptors; various receptors respond to discrete parts of an odor
structure; brain must determine the precise combination of activated receptors to distinguish a smell
Olfactory Receptors
humans 30 genes, some lost (converted to pseudogenes) during evolution
one of the largest G-protein receptors, linked to cAMP and IP3
part of 7TM receptor family (7 times passes membrane), odorant complemantarity determining regions in TM3, 4, 5
odorant selectivity – has hundreds of different genes for odorant receptors, only one in each OSN
G-protein Activation
Odorant + OR activates G-protein Effect
ATP into cAMP Ca influx = action potential Ca opens Cl
AC G-olf
channels electrical excitation
PLC Other G-protein production of IP3 + DAG act on ion channels and ↑ Ca levels
Ca and GC activating protein Other G-protein produce cGMP augments cAMP signal
G-olf G-protein in receptor cells, similar to Gs, made up of α active portion, and β & γ regulatory subunits, α-GDP is
inactive, α-GTP is active
α subunit has GTPase activity after activating AC = acts as regulatory mechanism
Cl influx to generate potential is unusual, OSN actively accumulates Cl against electrochemical gradient
Adaptation
Ca ions entering cyclic nucleotide-gated channels Ca acts with calmodulin ↓ cAMP sensitivity requires
stronger odor stimulus to produce sufficient cAMP
Acts via closing of Ca channels
Theories of Olfaction
Theory By Hypothesis
Molecular lock & key hypothesis of Moncrieff ability to bind depends on odorant shape and electrical charge
shape stereochemical theory of Amoore geometry of odorants = 7 primary odors
Davis and Taylor odorant molecules diffuses across membrane ion pore
Diffusion pore
formation
Rosenberg carotenoids + odorants = semiconductor current activate
neurons; Problems: receptors have no pigment and weakly
Piezo-effect odorous short chain alcohols gave greater increase in
semiconductor current than smellier long-chain alcohols.
Disorders
Disorder Notes
temporary or permanent loss of smell, most common; due to head trauma, upper
respiratory infection or nasal/sinus disease; may be due to smoking, crooked nose /
Anosmia / Anosphrasia
septum, medication,s drugs, aging, lead poisoning, insecticides, radiation therapy,
idiopathic loss
Hyposmia / Partial
diminished sense of smell, after acute influenza
Anosmia
Dysosmia distorted sense of smell, smells non-existent unpleasant odors
Hyperosmia increased sensitivity to smell, neurotics or histrionics
Kallman’s syndrome genetic, anosmia + ↓ functional activity of gonads
inability to distinguish a particular odor due to fatigue or desensitization, test a max of 3
Odor fatigue
odors when testing fragrances
Other notes
Astronauts = Capillary pressure = ↓ smell and taste
Women can differentiate HLA alleles via smell; ↑ HLA = preference
Amorphallus titanium – largest and smelliest (putrefying roadkill) flower, petals like rotting flesh, putrescin and
cadaverin
GnRH develop in olfactory tract
Vomeronasal Organ – bottom of nasal septum, active in animals, pheromone receptors, involuted in humans