Lecture 10 - Taste and Smell - Chemosensation Flashcards

1
Q

What is the main difference in chemosensation between animals, single cell prokaryotes and plants?

A

Single cell prokaryotes and plants have chemosensation but it is not neuronal based - organised by other more acient processes

SC prokaryotes - can orient towards and move up a gradient towards nutrients through chemical sensing
Plants: can orient towards air-borne chemicals allowing growth towards food sources

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2
Q

What actions do smell and taste guide?

A
  • food (nutritive value)
  • mate selection
  • danger
  • poison
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3
Q

What is they typical response to bitter/sour and sugary food?

A

Bitter/sour - rejection, bitter compounds often toxic

Sugars: appetitive, contain imp nutrients

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4
Q

What is the main difference between Olfaction and gustation?

A

Olfaction is the detection of chemicals at a distance and gustation requires direct contact with the relelvent chemical

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5
Q

What is the organisation of detection in olfaction and gustation?

A
  • Olfaction is optimised for combinatorial detection of vast numbers of odourants
  • Gustation is required to categorise tastants into defined non-overlapping modalities (bitter, sweet, sour, salty umami)
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6
Q

In insects and mammals, why are the major gustatory and olfactory organs closely associated?

A

may be due to evolutionary mechanisms

also functional - odour is used to attract but still assess before material is placed near the gustatory system

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7
Q

What is flavour a fusion of?

A
taste and odour
also texture (somatosensory)
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8
Q

Why does eating glucose result in more insulin production that direct injection into the brain?

A

Taste receptors are present in the gut in endocrine cells

  • cells produce hormones = incretins
  • incretins stimulate the secretion of insulin
  • this helps to prime the metabolic system and signals satiety in the brain when sufficient incretin is produced
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9
Q

What do taste receptors in the gut for ‘sour’ mediate?

A

a slowing down of stomach emptying

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10
Q

Where are taste receptors present?

A

Mouth
Gut
Brain

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11
Q

How do taste receptors in the brain react to a 24hr period of food deprivation?

A

Increase in T1R1 and T1R2 expression (involved in metabolic regulation)

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12
Q

What is throught to be a ‘taste’ receptor for fat?

A

Not entirely sure

-‘filliform’ papillae detect texture and fat may be partially a texture within food

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13
Q

What are our innnate respnses to sugar and tastants such as coffee?

A

Innate response to sugar is to eat a LOT of it, without any prior exposure
-On first exposure to coffee we will often reject it, although learn to like it on subsequent exposures

Taste is a driver of both innate and learnt behaviours

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14
Q

What are the functions of taste and what are the taste qualitites of bitter, sweet, sour, salty, and unami - what are these mediated by?

A

Function: prepare the gut for digestion (TIR and TIIR), and other organ function/activation
Bitter - poison? Inially reject e.g. coffee, learn to like
Sweet - sugar, carbohydrate (energy)
Sour - organic acids
Salty - Sodium (ion channel transduction)
Umami - L-amino acids, nucleotides

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15
Q

What is the average lifespan of taste receptors and how might this contribute to loss of taste with age?

A
  • taste receptors are constantly renewed throughout life

- lifespan of two weeks

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16
Q

What are the structures of taste buds, where are these found on the tounge and what do they all contain?

A
3 structures in the tongue
Circumvallate
Foliate
Fungiform 
All contain taste buds which contain taste receptors to allow solubalised (by saliva) taste molecules through 
Circumvillate 
-taste buds on side of the pore
-located at the back of the tongue
Foliate 
-taste buds on side of the pore
-located at the middle edges of the tongue
Fungiform 
-taste bud on top of the pore
Located at the front of the tongue
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17
Q

What are the different receptors for sweet, umami, sour and bitter - how was this discovered experimantlly?

A

T1R receptors: GPCRs -sweet and umami
T1R1+T1R3 = umami
T1R2+T1R3 = sweet
T1R3 is the common receptor

T2R receptors:GPCRs bitter

PKD1L3+PKD2L1: TRP receptors - sour

Experimentally
Recordings from single afferent fibres in receptor KO mice

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18
Q

T1Rs are a major difference in taste preference between mammals - what does this suggest about these G protein coupled receptors?

A
  • appear to evolve rapidly
  • v large sequence differences between mouse and human in the extracellular sequence
  • reflects differences in diet and niche
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19
Q

What are the features of T1Rs and T2Rs?

A
  • T1Rs - low affinity, so saturation not easily reached for an appetitive stimulus
  • T2Rs - higher affinity, necessary to detech noxious foodstuffs more rapidly and at lower concentrations to generate a safety margin
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20
Q

What does the taste of saccharine elicit at different concentrations?

A

Elicits sweet taste at lowconcentrations, sour at high conc

-may be due to the broad tuning of both sweet and sour receptors and the differing affintiy of the ligand

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21
Q

How is the action of PKD2L1 a clear indicator of the labelled line theory?

A

Knocking out neurons expressing this channel abolishes the ability to taste ‘sour’ without affecting other tastes

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22
Q

What experiment was done to show strong evidence for the labelled line model for tastants?

A
  • took sweet promotor and conjugated to the bitter receptor so that the bitter receptor was expressed in the sweet cell
  • generated at attractive response to a bitter tastant that would normally be shown in response to a sweet tastant
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23
Q

What are the features of taste buds~?

A
  • up to 100 polarised neuroepithelial cells which form columnar islands in the oral cavity
  • have a neuronal and glial characterisitics
  • tips directly contact apical surface - cell exposed to damage therefore continuously renewing
  • although there is some regional sensitivity there is no taste map
  • Types I, II, III, IV (Type I-III imp)
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24
Q

What techniques can be used to show the localisation of the different types of tastebuds?

A
  • electromicrograph
  • fluoresence labelling with GFP
  • immunostaining
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25
Q

What are the three major classes of taste cells?

A

Type I glial-like cell
Type II receptor cell
Type III presynaptic cell

26
Q

What are the features of the Type I glial-like cell?

A

Type I glial-like cell

  • degrade or absorb neurotransmitters
  • clear extracellular K+ that accumulates after action potentials in receptor and presynaptic cells
  • K+ may be extruded via an apical K channel e.g. ROMK
27
Q

What are the features of the Type II receptor cell?

A

Type II receptor cell

  • sweet, bitter and umami taste transduction, ATP then released through pannexin1 (Panx1) hemichannels
  • extracellular ATP then excites ATP receptors (P2X, P2Y) on sensory nerve fibres and on taste cells
  • transmits signal to nerve with no synaptic vesicles
28
Q

What are the features of the Type III presynaptic cell?

A

Type II presynaptic cell

  • contains surface glycoproteins, ion channels
  • inv. in neurotransmitter synthesis, excitation and transmitter release
  • release serotonin (5-HT) which inhibits receptor cells
  • sour stimuli (and carbonation) directly activate presynaptic cells
  • only cells to form identifieable synapses with nerves
29
Q

What is the mechanism by which taste quality is transduced in receptor Type II cells?

A

1-sweet, bitter and umani ligands bind taste GPCRs
2- activate a phosphoinositide pathway that through IP3 and the IP3R3 calcium channel, elevates cytoplasmic Ca2+ concentrations and depolarises the membrane via the TrpM5 cation channel
3- the combined action of the elevated Ca2+ and membrane depolarisation opens the large pores of gap junction hemichannels (likely composed of Panx1) resulting in a ATP release

30
Q

What is the mechanism by which taste quality is transduced in pre synaptic Type III cells?

A

1-IN presynaptic type III cells, organic acids (HAc) permeate through the plasma membrane and acidify the cytosol
2-intracellular H+ is believed to block a proton-sensitive K channel and polarise the membrane
3- voltage gated Ca2+ channels would then elevate the cytoplasmic Ca2+ to trigger exocytosis of synaptic vesicles

31
Q

How is the salty taste of Na+ detected?

A

By direct permeation of Na+ ions through membrane ion channels including ENaC to deparise the membrane

32
Q

What are the two current models of taste perception?

A
  • labelled line

- Across fibre models

33
Q

which of the two current models of taste perception are favoured and how does this model indicate saliency?

A

Labelled line model

Cross talk between types of cells to indicate saliency

34
Q

What is the mammalian olfactory system?

A

chemosensation at a distance, closely linked with the respiratory and gustatory appartus and aided by turbulent air editties
-water soluble molecules bind receptors

35
Q

What is the mammalian olfactory epithelia?

A
  • lines the nasal cavity and allows direct access to oderant molecules
  • has layer of protecting mucus
  • neurons are turned over every ~90 days
36
Q

What are the levels of olfactory receptors in humans and mice and what is their main feature?

A

OR human: 950
OR mouse: 1500

Each olfactory sensory neuron expresses only one type of olfactory receptor

37
Q

What is the anatomy of the rodent peripheral olfactory system?

A
  • expression of only one receptor per OSN generates a topographic map
  • 5-10million olfactory sensory neurons in the olfactory epithelium
  • 1800 glomeruli in each olfactory bulb
  • 103 convergence of primary sensory axons onto each olfactory glomerus (lies at the heart of coding)
  • also have vomeronasal tract for non volites (pheromones)
38
Q

What are the structures involved in the mammilain olfactory system?

A

olfactory epithelium
turbinate bones
olfactory bulb
-cribiform plate of ethmoid

39
Q

What are the features of signal transmduction in the OSN and the organisation of coding?

A
  • each OSN expressed just a single allele of a specific OR (one receptor-one neuron rule)
  • neurons expressing one OR may be distributed broadly in the olfactory epithelium but send projections to discrete glomeri in a spacially invariant pattern
  • mirror patterning will innervate one glomerus in medial hemisphere and one in the lateral hemisphere of the olfactory bulb
40
Q

What is the process of signal transmduction in the OSN?

A

1-binding of an odourant to its cognate OR results in activation of heterotrimeric G protein (Gαolf plus Gβγ)
2-activatied Gαolf activates type III adenylcyclase (AC3) leading to the production of cyclic AMP from ATP
3-cAMP gates the cyclic nucleotide-gated (CNG) ion channel, leading to an influx of Na+ and Ca2+ depolarising the cell
4- initial depolarisation is amplified through the activation of Ca2+ dependent Cl-channel
5-cAMP also activates protein kinase A (pKA) which can regulate other intreacellular events, including transcription of cAMP-regulated genes

-OR-mediated activity also leads to the transcriptional regulation of cAMP response element binding protein (CREB)-dependent gene expression via CREBs phosphorylation via PKA

41
Q

How is selection of a particular OR gene by the cell thought to occur?

A

via interaction of a cis-regulatory locus control region with the proximal promotor of a single OR gene within a cluster of OR genes
-choice is stabilised and the expression from all the other OR genes is silences by an OR dependent feedback loop which ensures the expression of a single OR per sensory neuron

42
Q

What is the combinatorial coding of olfactory information?

A

Patterns of receptors are activated by an odour, each to a different magnitude

  • the pattern of receptor activation elicited by a particular compound is thought to represent that chemical’s chemical identity, and discrimination and appropriate responses occur at higher order structures such as the piriform complex
  • some receptors are tuned finely, others broadly
43
Q

What is zone-to-zone projection of OSN axons to the olfactory bulb along the dorsal vental axis shaped by?

A

by complementary gradients of the chemorepellent molecules Slit1 and Sema3F and their receptors Robo2 and Nrp2 respectively

44
Q

What is innervation of the lateral olfactory bulb dependent upon?

A

IGF signalling

-which may function to counteract a default tendancy of all olfactory neurons to project medially

45
Q

What is the projection of olfactory sensory axons along the olfactory bulbs anterior-posterior axis dependent upon?

A

the level of intracellular cAMP

  • which inturn regulates the expression of the acon guidance receptor Nrp1
  • by modulating the expression levels of axon guidance receptors such as Nrp1 the sensory axons are either more or less sensitive to guidance cues found in the OB or along the projection pathway
46
Q

What are the main features of ORs?

A
  • largest gene family in the mammalian genome
  • expressed in OSN
  • one receptor per neuron
  • receptor determines axon guidance and generation of a topographic map
  • receptors can be broadly and narrow tuned
47
Q

How much more sensitive is the insect olfactory system to the mammalian system?

A

2/3 times more sensitive

48
Q

What occurs when flies walk on a substrate with an appetitive content?

A
  • chemosensors on their feet generate a ‘Proboscis extension reflex’
  • so flies detect food as they walk on it allowing an extension of the mouthparts to the substrate
49
Q

Where are flies chemosensory senses organised?

A

Mostly sensilla on their antenna - particularly the 3rd antenally segment

  • smaller strcuture at the top of the proboscis (maxillary palp) also involved
  • have chemosensitive receptors on many parts of the body, wing, abodomen, feet
50
Q

What are sensilla?

A

sensory projections on the Antenna and maxillary palp

51
Q

What are sensilla divided into?

A

Large and Small Basiconic and Coeloconic

-classification structural and anatomical

52
Q

How do sensilla differ in males and females (flies)?

A

419 in males
~457 in females

may be involved in courtship

53
Q

What are the structures of the proboscis?

A

DCSO - Dorsal Ciberial Sense Organ
VCSO - Ventral Cibrarial Sense organ
LSO - Lateral sense organ

54
Q

What is the organisation of olfactory sensation in flies in the antenna?

A
  • each olfactory receptor is expressed in a subset of sensilla, except Or83b which is expressed in all sensilla
  • olfactory neurons from similar anatomic regions project from the antenna through the suboesophageal ganglian (SOG) to the antennal lobe
  • the antennal lobe sends projection neurons (dendrites=ganglions) to the mushroom body, specifically to the mushroom body calyx (MBC) which is a large tripartite structure that is known to mediate olfactory memory
55
Q

What are the features of Drosophila olfactory receptors and gustatory receptors?

A
  • may be G protein coupled
  • 7 transmembrane domains
  • inverse conformation to mammalian ORs
  • Oligate heterodimer
56
Q

What are the types of sensory structures in antenna, palp and larvae?

A
Antenna
Basiconic sensilla
-Food odours and CO2
Trichoid sensilla
-Pheromones
Coeloconic sensilla 
-Food odours, water vapour, ammonia, putrescine 

Palp
Basiconic sensilla
-food odours

Larva
Dorsal organ dome
-food odours

57
Q

How many ORs are there in drosophila/mosquito/honeybee?

A
Drosophila
62
Mosquito
79
Honeybee
157
58
Q

What genes involved in chemosensation lead to anosmic drosophila mutants?

A

Or83b/ORCO - necessary for smell

59
Q

Give an example of how a single pheremone can elicit different behaviours in the two fly sexes

A

1- 11, cis-vacceyl-acetate is exued by male flies and causes agression behaviour
2-loss of OR67d causes males to inappropriately court other males
3- the glomerus that OR67d projects onto expresses the TF fruitless, critical for generating divergent sexual behaviour between males and females
-females mutant for OR67d however are not as receptive to courting males

60
Q

What are the olfactory receptor neuron, primary centre and secondary centres of mammals and insects?

A
Olfactory receptor neuron 
M: olfactory epithelium
I: Antenna and Maxillary Palp
Primary centre
M: Olfactory bulb 
I: Antennal lobe
Secondary centres
M: Piriform cortext, Olfactory tubercle, Entorhinal cortext
I:MB calyx, Lateral horn