Chemoreceptors

- Detect smell, taste, chemicals in blood for food, sexual males, predators, harmful
substance, hypoxia
Nasal conchae
- Long, narrow, curled shelves of bone that protrudes into the breathing passage
- Scroll-like bones, force air into the passages between where temperature and
humidity can be controlled
Odorants
- Ester, terpenes, lactones, ketones, aldehydes, thiols
Epithelium
- Pseudostratified epithelium is comprised of a mono layer of cells
- Nuclei positioned to look stratified
- Usually considered as pseudostratified columnar epithelium
Olfactory epithelium
- Yellow patch, site of olfaction
- Pseudostratified epithelium
- Roof of the nasal cavity
- Problem: ill positioning, solution: sniffing

- Nuclei of the sustentacular cells tend to be in the top portion of the layer

- Nuclei of the olfactory receptor cells tend to be in the middle portion

- Nuclei of the basal epithelial cells tend to be in the bottom portion.

- Epithelium contains olfactory neurons

- Epithelial cells are yellowish (lipofuscin)

- At the base of the olfactory epithelium lie the olfactory stem cells (basal cells)

- Olfactory nerve (CN I) penetrates cribriform plate

- Spherical structures of where mitral cells and olfactory cells synapse: Glomeruli
(1800)

- Mitral cells are output cells of the olfactory bulb

Bowman’s glands
- Bowman's glands are situated in the olfactory mucosa
- Beneath the olfactory epithelium
- In the lamina propria
- Secrete the gel-forming mucin
- Secrete proteins lysozyme, amylase and IgA
- Some evidence of production of odorant binding protein
Olfactory neurons
- Olfactory neurons are bipolar neurons
- The apical dendrite ends in many cilia
 - Cilia protrude from the olfactory receptor into the mucus
- Surface of cilia is covered with olfactory receptors
- Olfactory receptors are G protein-coupled receptors
Olfactory receptors
- Each receptor binds to a variety of odour molecules, affinity can vary
- Affinity difference causes differences in activation patterns resulting in unique
odorant profiles.
- Activated OR activates the intracellular G-protein, adenylyl cyclase (AC) and cyclic
AMP (cAMP)
- cAMP opens ion channels in the cell membrane
- Na+ and Ca++ enter cell, Cl- out
- Depolarization, AP
Olfactory pathway
- Same receptor = same glomerulus
- Each odour activates a certain set of glomeruli
- Olfactory bulb also contains amacrine cells
- Axons of olfactory neurons synapse in the olfactory bulbs
- Synapse with mitral cells (2nd order neurons) in glomeruli
- Same receptor = same glomerulus
- Each odour activates a certain set of glomeruli
- Olfactory bulb also contains amacrine cells
- From olfactory bulb, signal goes via olfactory tract (mitral cell axons) to the olfactory
cortex
- Some to the frontal lobe: conscious interpretation
- Some via hypothalamus and amygdala (limbic system): emotional response
Peripheral stem cells in the olfactory system
- Olfactory receptor cells are constantly turned over.
- The source is stem cells within the olfactory epithelium.
- This is a highly regulated process and is being used as a model of neuronal stem cell
biology.
- The axons of new ORNs penetrate the olfactory bulb.
- Special glial cells (ensheathing) facilitate this; ordinary adult glia block axonal
regeneration; so the ensheathing cells are of interest to molecular neuroscientists
interested in axonal regeneration.
- Further, the new ORN axons make correct connections in the OB: that is, to the
glomerulus specified by the receptor type they express. The mechanism for such
specific regeneration is unknown and of intense interest.
Amoore’s theory
- Problems with Amoore’s Theory
– HCN smells the same as benzaldehyde – ‘almonds’ - 350 different olfactory receptors in
humans
- Problem with all theories so far:
350 receptors in mammals, if each responds to a few odorants then we should be able to
smell ~1000 odours
The range of odours we can detect is at least 10 000

Labelled line theory

 - Each of the 350 receptors sends a ‘labelled-line’ to the brain
- Enables brain to determine which of the 350 receptor types was stimulated and so
determine odorant • Would limit us to about 1000 detectable odorants only!
- This theory must be discarded!
Combinatorial odour code theory
- Since each receptor responds to a small range of different odorants:
- Brain can distinguish odours based on the pattern of receptor activation – i.e. which
of the 350 receptors activate gives a unique combination identifying each odour
- This allows an almost infinite number of odours to be detected, though we only
discriminate about 3000 of these
Intrinsic signal imaging
- Image olfactory bulb under red light (630 nm)
- Visualise flow of oxygenated blood in olfactory bulb
- Only those glomeruli activated by an odour use a burst of oxygen
- Each odour then gives a distinct pattern of olfactory bulb (glomeruli) activation
- Individual odorant may activate many glomeruli
- An individual glomerulus may be activated by many different odours
- Pattern of glomeruli activated characterises odorant
Olfactory adaptation
- Adaptation: The decrease in sensitivity of a sensory system over time when a
continuous non-changing stimulus or a rapidly repeating stimulus is applied.
- Short-term adaptation is due to the refractory period during which calcium levels
inside cell are returned to normal (calcium → negative feedback)
Perceiving complex odours
- Some of the odorant molecules found in Jasminum officinale (jasmine):
- In mixtures of 2 – 4 odorants people can distinguish individual component odorants
- Power of discrimination is lost for more complex mixtures
- Configurational hypothesis – the component odours form a recognisable signature
which is perceived as a single distinctive odour
- HCN and benzaldehyde – very different structures - both smell of almonds, both
produced by almond!
Olfactory system: olfactory bulb
- It appears likely that the code for odorant identity is spatio-temporal:
- An odorant will activate different but overlapping populations of OB neurons and the
activated cells will have different patterns of spiking discharge.
Chemoreception
- Olfaction involves two cranial nerves:
- The olfactory nerve is responsible for most of our sense of olfaction
- The olfactory nerve ends in the olfactory bulb. Modified epithelial cells (a type of
bipolar neuron) lining the nasal cavity below the olfactory bulbs send their axons
through pores in the cribriform plate to synapse in the olfactory bulb.
- But branches of the trigeminal in the nasal cavity are responsible for detecting the
smell of burning.
Gustation
- Epiglottis folds over windpipe when swallowing. However, humans can breathe and
speak whilst chewing, raising the risk of choking on one’s food!
- Pharynx is divided into nasopharynx, oropharynx and laryngopharynx.
 - Laryngopharynx – extends from base of skull to top of soft palate.
- Uvula – the ‘dangly ball’ in the back of the throat – involved in speech and gag reflex.
- Pharyngotympanic tube (eustacian tube) connects throat to inner ear to equilibrate
the pressure on both sides of the ear drum.
- Oropharynx (oral end of pharynx) extends from hyoid bone level to uvula.
Receptor organs are taste buds
- Found on the tongue
- On the tops of fungiform papillae
- On the side walls of foliate papillae and circumvalite papillae
- Also found on palate
Papillae
- Foliate papillae: laterally located, lost early.
- Fungiform papillae: mushroom shaped.
- Filiform papillae: non-tasting. Mechanical friction.
- Circumvallate papillae: largest. Inverted v – each houses 100’s of individuals buds.
Structure of a taste bud
- Small neurons called gustatory hairs. Flask shaped, 50–100 epithelial cells:
- Basal cells—dynamic stem cells ➢ Gustatory cells—taste cells
- Microvilli (gustatory hairs) project through a taste pore to the surface of the
epithelium
Taste sensation
- There are five basic taste sensations
- 1. Sweet — sugars, saccharin, alcohol, and some amino acids
- 2. Sour—acids + hydrogen ions
- 3. Salt — metal ions, NaCl; KCl tastes bitter
- 4. Bitter — alkaloids such as quinine and nicotine
- 5. Umami—amino acids glutamate and aspartate
Taste transduction
- In order to be tasted, a chemical: Must be dissolved in saliva, Must contact gustatory
hairs
- Binding of the food chemical (tastant): Depolarizes the taste cell membrane, causing
release of neurotransmitter, Initiates a generator potential that elicits an action
potential
- The stimulus energy of taste causes gustatory cell depolarization by: Na+ influx in
salty tastes (directly causes depolarization), H+ in sour tastes (intracellular pH !), Both
un-dissociated acid molecules, e,g, CH3COOH, and H+ contribute to sour taste – it
appears to be the intracellular pH of the taste receptor that is measured! Mechanism
of how the acids effect intracellular pH not clear. GPCR → G protein gustducin in
sweet, bitter, and umami tastes (leads to release of Ca2+ from intracellular stores,
which causes opening of cation channels in the plasma membrane)
Gustatory pathway
- Cranial nerves VII, IX, X carry impulses from taste buds to the solitary nucleus of the
medulla
- Impulses then travel to the thalamus (in medial lemniscus) and from there fibers
branch to the: Gustatory cortex in the insula, Hypothalamus and limbic system
(appreciation of taste)
 - Facial nerve: fungiform papillae

- Glossopharyngeal nerve: circumvallate and foliate papillae

- Vagus: epiglottis, larynx, soft palate

- Trigeminal: somatosensory

Tastant mixtures

- Tastants of same quality may enhance each other so the overall flavour is stronger
than expected:

- Mixtures may have reduced taste

- One tastant may mask another