Lecture 2 - smell And Taste Flashcards
What are the traditional senses?
Smell (olfaction) Taste (gustatory, gustaoception) Hearing (audioception) Sight (vision, opthalmoception) Touch (tactioception)
Temperature
Thermoception
Kinesthetic sense
Propioception
Pain
Nociception
Balance
Equilibrioception
Various internal stimuli
Chemoreceptors
What is the cranial nerve for smell?
Olfactory (I)
What is the cranial nerve for sight?
Optic (II)
What is the cranial nerve for hearing?
Vestibulocochlear (VIII)
What is the cranial nerve for Taste?
Glossopharyngeal (VII, IX, X)
What is the cranial nerve for touch?
Trigeminal (V)
Where are the most common kind of receptor properties found?
Terminal ramifications of a sensory axon that belongs to a pseudo-unipolar ganglion cell
Where are the receptors found in the sense organs located?
Sensory cells that transmit a signal to a ganglion cell
What are the perception of odours closely associated with,
Memories
Emotions
Moods
For humans, what important role does sense of smell have?
Interpersonal communication
Humans can sense over how many different odours?
10,000
What is the detection threshold?
As low as parts per million or even billion
For humans what is smell?
Largely an aesthetic sense
How sensitive is smell in scent-tracking in many animals?
~ 10.000.000
What does smell direct animals to?
Food
Potential mates
Away from danger
Where does the olfactory signalling in humans start?
Nasal cavity - olfactory epithelium
Where does the olfactory receptor neutrons send their axons to?
Olfactory bulb
Where does the second order Neuton transmit to?
Higher cortical areas
What is accessory olfactory pathway (AOP) used for?
Pheromone defection
What is pheromone ?
Species specific
Affect animal behaviour
What is MOP defined as?
Main olfactory system (MOS)
What is AOP defined as?
Accessory olfactory system (AOS)
What does the large family of olfactory receptors recognise ?
Numerous ligands
How many olfactory receptors are there?
~1000
What are olfactory receptors?
GPCRs belonging to the odorant receptor gene family
Class “A” GPCR (Opsin or B-adrenergic receptors)
Who was OR discovered by?
R.Axel
L.Buck
What does a single odour activate?
Multiple ORs
Different combination of ORs are needed for odour recognition
How many OR genes in mouse?
1300 (5% of genome)
How many ORs in humans?
500 ORs (2% of genome. 50-95% sequence homologous)
How many ORs in Zebrafish?
100
How many ORS in C.elegans?
1500 (7% of the genome)
What does C.Elegans consist of?
Nematode
Roundworms
How many neutrons and chemoreceptor genes does C.elegans have?
302 neutrons
1500 chemoreceptor genes (7% of the genome)
How many sensory neutrons express most of the receptors?
32
In mammals, how many olfactory nephron expresses one OR gene?
1
What are olfactory receptor neurons?
Bipolar cells within olfactory epithelium
What does ciliated endings contain?
Odorant receptors that detect odours
Where does axons project to?
Glomeruli in the olfactory bulb
Where does mitral/tufted cells send information to?
Higher brain centres
What is the mechanism of odorant receptors?
- Odorants bind to receptors
- Olfactory receptor cells are activated and send electrical signal
- The signals are related in glomeruli
- The signals are transmitted to higher regions of brain
What does the sensory transduction of odour molecules involve?
Cylic AMP second messenger pathway
What does activation of a receptor by odour molecules activate?
GTP binding protein
What does GTP-binding protein activate?
Adenylate cyclase which produces cyclic AMP
What does cAMP activate?
Cationic cyclic nucleotide-gated channel
Chloride channel
What are olfactory cells?
First order neurons
What are mitral cells?
Second order neurons onto which axons of olfactory cells synapse
What is the neurotransmitter between olfactory cells and mitral cells?
Glutamate
Where does the olfactory neurons send axons to?
Olfactory bulb
What is olfactory bulb composed of?
Layers
What is the peripheral layer?
Glomerular later where the axons of the olfactory neurons terminate
What do many axons converge into?
Single glomerulus
In rats, how many olfactory neurons converge onto ~ 1500 glomeruli?
~ 15 million
What do neurons with different ORs send axons to?
Different glomeruli
What does activation of different receptors lead to?
Activation of different glomeruli
What does combination of activated glomeruli encode?
Smell
What do olfactory neurons with the same receptor project to?
Same glomerulus
What do mitral cells synapse onto?
Single glomerulus
What are periglomerular cells?
Inhibitory interneurons that synapse within and and between glomeruli
What are granule cells?
Inhibitory interneurons that synapse between mitral/tufted cells
What do inhibitory cells use?
GABA/Dopamine/Glycine
Mitral cells
Project directly into olfactory cortex (Glutamate)
What is the main cortical area receiving olfactory input ?
Pyriform cortex
Where does the conscious perception of smell happen?
Orbitofrontal cortex
Where does pyriform cortex project to?
Thalamus via orbitofrontal cortex
Where does olfactory tubercle (OT) receive input from?
Tufted cells - project to hypothalamus
Cortico-medial group amygdala (AMYG)
Input from accessory olfactory bulb - hypoth
Transitional entorhinal cortex (TEC)
Information reaches hippocampus
Anterior olfactory nucleus (AON)
Cells just posterior to olfactory bulb
What does NHLDB stand for?
Nucleus of horizontal limb of diagonal band
What does MD stand for?
Mediodorsal
What does VNO stand for?
Vomeronasal Organ
Where is the primary olfactory cortex located?
Uncus
Where does POC project to?
Medial thalamus - orbitofrontal cortex
Where does olfactory tubercle project to?
Hypothalamus
What does pheromone provide information about?
Individuals social, sexual or reproductive status
What does female snake produce?
Substance which can induce courting behaviour in male snakes
What does slave maker and emit a blend of?
Chemicals that signal alarm
What can male dogs sniff?
Female in heat
Where are pheromones detected in?
Accessory olfactory pathway (AOP)
Where are pheromone detector barons located in?
Vomeronasal Organ (VNO) - anterior nasal septum
Where can detection can happen?
Openings connecting VNO to either oral or nasal cavities
Where does the axons from sensory neurons in VNO travel to?
Accessory olfactory bulb (AOB)
Does AOP and MOP overlap?
No - functional segregation
What is the sensory epithelium in VNO divides into?
Apical and Basal layer
What does neurons in apical layer express?
V1R receptors
What do neurons in basal layer express?
V2R receptors
Where does the apical layer project to?
Anterior AOB
Where does the basal neurons project to?
Posterior AOB
What does mitral cells interconnect?
Different glomeruli
What does binding of pheromones to V-R receptors in microvilli activate?
PLC through the G protein
What does PLC produce?
Second messengers DAG and IP3
What does the gating of TRPC3 channel by DAG allowable
Depolarising influx of Na+ and Ca2+
What can intercellular calcium gate?
A ca2+ Activated Cl- current
How many functional VR1 genes in mice?
165
How many functional VR1 genes in rats?
106
How many functional VR1 genes in humans?
2
How many VR1 genes in humans are pseudo genes?
115/117
How many VR2 genes in mice and rats?
~60
Where do you find a lack of VR2 genes in?
Dogs
Cows
Primates
What is a TRPC2 gene?
Non-functional pseudogene in humans
What are other factors that contribute to taste?
Thermoreceptors Mechanoreceptors - texture Nociceptors Sight Olfaction
What is the gustatory system?
Detect soluble chemicals at a short range
What is the primary purpose of gustatory system?
Identify sources of nutrient rich food and to avoid toxins
What is the amino acid and sugars detect threshold?
10-100 mM
What is bitter detection threshold?
< 0.02 mM
What are the taste modalities in mammals?
Sweet Bitter Sour Salty Umami
Where does Mammal taste recognition start?
Tongue where taste buds are located
What does taste buds contain?
Taste deception cells (~100) - epithelial on nature
depolarise and release neurotransmitter
Apocalypse, what are receptor cells covered by?
Microvilli
Which contain receptors for recognition of taste
Basally, what does the cell synapse with?
Gustatory nerve
What is the neurotransmitter binding to purinoreceptors?
ATP
What are the other location for taste receptors?
Epiglottis
Soft palette
What occurs in the gustatory cortex?
Gustatory sensory integration occurs
What does the gustatory cortex consist of?
Insular cortex
Posterior limb
Internal capsule
Frontal operculum
What are the main nuclei in rodents of taste perception?
NST - nucleus of solitary tract
PBN - parabrachial nuclei
What is the main nuclei involved in humans for taste perception?
NST - nucleus of solitary tract
VPMpc - ventral posteriomedial nucleus
Where are various taste bud population found In?
Fungiform papillae - anterior tongue
Palate and foliate papillae - posterior tongue
What are taste buds innervated by?
Branches of VIIth, IXth, Xth cranial nerves
What does the buds on palate and epiglottis contact?
Xth cranial nerve
What is the gustatory pathway in humans?
The afferent fibres collect in rostral solitary nucleus of Medulla oblongata
The axon ascend ipsilaterally into central tegmental tract of brain stem terminating in parvocellular division of central posteriomedial nucleus in thalamus
Go into gustatory neocortex
Enter limbic system
What is the gustatory pathway in rodents?
The axon of the peripheral gustatory fibre in facial, glossopharnygeal and vagal cranial nerve lead into nucleus of solitary tract in Medulla oblongata
Leads to parabrachial nucleus followed by ventral posteriomedial nucleus in thalamus
From thalamus the neurons project into gustatory neocortex
Project into hypothalamus and amygadala
How many taste cells does the taste bf contain?
50-100 taste cells
What does epithelial receptor cells make synaptic contact with?
Dendrites of cranial nerve VII, IX or X
Cell bodies lie within cranial nerve Ganglia
How is the characteristic spindle shape of taste receptor cells revealed?
Subset of taste cells labelled with antibody against alpha-gustducin
Mice lacking T1R1
Do not detect amino acids
Mice lacking T1R2
Do not detect sugars
How are tests confirmed?
Behavioural testing of mice preference to different foods
What can humans detect?
Glutamate
What can humans detect?
Sweetness in aspartame
What does the activation of a receptor activate?
Heterotrimeric G protein
Activates phospholipase C-B2 (PLC B2)
Produce secondary messenger IP3 and DAG
Activation of PLC-B2 lead to opening of TRPM5 ion channel and cell depolarisation.
What are sour taste associated with?
PKD1L3/PKD2I1 channels
Intensity of sour taste various with proton concentration
What are salty taste mediated by?
ENaC sodium channels
Expressed apically in salt sensing cells
Na+ diffuses passively to cause depolarisation
Removed at the vascular membrane NaK-ATPase
What are 75% of humans able to detect?
Phenylthiocarbamide
Found in Brussels sprouts
What allows for full taste perception?
Cingulate gyrus
Secondary neocortical taste area in caudolatetal orbitofrontal cortex