Lecture 12: olfaction: the insect antennal lobe References: H C Mulvad, thesis (http://www.nordita.dk/~mulvad/Thesis), Ch 2http://www.nordita.dk/~mulvad/Thesis.

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Presentation transcript:

Lecture 12: olfaction: the insect antennal lobe References: H C Mulvad, thesis ( Ch 2http:// G Laurent, Trends Neurosci (1996) M Bazhenov et al, Neuron and (2001) Dayan & Abbott, Sect 7.5

Olfaction (smell)

The oldest sense (even bacteria do it)

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar)

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy:

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex ~ receptor cells, several hundred types (distinguished by receptor proteins)

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex ~ receptor cells, several hundred types (distinguished by receptor proteins) any cell responsive to a range of odorants:

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex ~ receptor cells, several hundred types (distinguished by receptor proteins) any cell responsive to a range of odorants: => an odor produces a characteristic pattern of activity across the receptor cell population

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex ~ receptor cells, several hundred types (distinguished by receptor proteins) any cell responsive to a range of odorants: => an odor produces a characteristic pattern of activity across the receptor cell population Receptor physiology: Receptor proteins (1 kind/cell): metabotropic, G-protein coupled, lead to opening of Na channels

Olfaction (smell) The oldest sense (even bacteria do it) Highly conserved in evolution (mammals and insects similar) Basic anatomy: Insects:receptor cells -> antennal lobe -> mushroom bodies Mammals: receptor cells -> olfactory bulb -> olfactory cortex ~ receptor cells, several hundred types (distinguished by receptor proteins) any cell responsive to a range of odorants: => an odor produces a characteristic pattern of activity across the receptor cell population Receptor physiology: Receptor proteins (1 kind/cell): metabotropic, G-protein coupled, lead to opening of Na channels, similar to phototransduction in retina

Antennal lobe ~ neurons in locust: 1130: 830 excitatory, 300 inhibitory in honeybee: 800 excitatory, 4000 inhibitory

Antennal lobe ~ neurons in locust: 1130: 830 excitatory, 300 inhibitory in honeybee: 800 excitatory, 4000 inhibitory Organized into glomeruli (bunches of synapes) (~1000 in locust, 160 in bee)

Antennal lobe ~ neurons in locust: 1130: 830 excitatory, 300 inhibitory in honeybee: 800 excitatory, 4000 inhibitory Organized into glomeruli (bunches of synapes) (~1000 in locust, 160 in bee)

Antennal lobe ~ neurons in locust: 1130: 830 excitatory, 300 inhibitory in honeybee: 800 excitatory, 4000 inhibitory Organized into glomeruli (bunches of synapes) (~1000 in locust, 160 in bee) Connections between AL neurons: dendrodentritic

Excitatory cells (PN) PN = projection neuron: axon takes its spikes out of the antennal lobe, to the mushroom bodies (+ other higher areas)

Excitatory cells (PN) PN = projection neuron: axon takes its spikes out of the antennal lobe, to the mushroom bodies (+ other higher areas) transmitter: ACh

Excitatory cells (PN) PN = projection neuron: axon takes its spikes out of the antennal lobe, to the mushroom bodies (+ other higher areas) transmitter: ACh

Excitatory cells (PN) PN = projection neuron: axon takes its spikes out of the antennal lobe, to the mushroom bodies (+ other higher areas) transmitter: ACh Dendrites have postsynaptic terminals in 1 or more glomeruli (10-20 in locust)

Inhibitory cells (LN) LN = local neuron: projects only within the antennal lobe

Inhibitory cells (LN) LN = local neuron: projects only within the antennal lobe no Na spikes, only Ca “spikelets”

Inhibitory cells (LN) LN = local neuron: projects only within the antennal lobe no Na spikes, only Ca “spikelets” transmitter: GABA

Inhibitory cells (LN) LN = local neuron: projects only within the antennal lobe no Na spikes, only Ca “spikelets” transmitter: GABA

Inhibitory cells (LN) LN = local neuron: projects only within the antennal lobe no Na spikes, only Ca “spikelets” transmitter: GABA Dendrites with postsynaptic terminals in several or all glomeruli

Antennal lobe responses:temporally modulated oscillatory activity patterns

~20 hz oscillations:

Antennal lobe responses:temporally modulated oscillatory activity patterns (No oscillations in input from receptor cells) ~20 hz oscillations:

Oscillations and transient synchronization membrane potentials

Oscillations and transient synchronization membrane potentials Local field potential In mushroom body: Measures average AL activity (cell in mushroom body)

Oscillations and transient synchronization membrane potentials Local field potential In mushroom body: Measures average AL activity PN firing transiently synchronized to LFP (cell in mushroom body)

Model (Bazhenov et al) 90 PNs, 30 LNs

Model (Bazhenov et al) 90 PNs, 30 LNs Single-compartment, conductance-based neurons

Model (Bazhenov et al) 90 PNs, 30 LNs Single-compartment, conductance-based neurons (post)synaptic kinetics

Model (Bazhenov et al) 90 PNs, 30 LNs Single-compartment, conductance-based neurons (post)synaptic kinetics Fast excitation, fast and slow inhibition

Model (Bazhenov et al) 90 PNs, 30 LNs Single-compartment, conductance-based neurons (post)synaptic kinetics Fast excitation, fast and slow inhibition 50% connectivity, random

Model (Bazhenov et al) 90 PNs, 30 LNs Single-compartment, conductance-based neurons (post)synaptic kinetics Fast excitation, fast and slow inhibition 50% connectivity, random Stimuli: 1-s current pulse inputs to randomly-chosen 33% of neurons

Bazhenov network

Excitatory neurons

Active currents:

Excitatory neurons Active currents: Na

Excitatory neurons Active currents: Na K

Excitatory neurons Active currents: Na K A-current

Excitatory neurons Active currents: Na K A-current Synaptic input

Excitatory neurons Active currents: Na K A-current Synaptic input Fast (ionotropic) synaptic currents (nACh and GABA A ): ( [O] is open fraction)

Excitatory neurons Active currents: Na K A-current Synaptic input Fast (ionotropic) synaptic currents (nACh and GABA A ): ( [O] is open fraction) [T] is transmitter concentration:

Excitatory neurons Active currents: Na K A-current Synaptic input Fast (ionotropic) synaptic currents (nACh and GABA A ): ( [O] is open fraction) [T] is transmitter concentration: exc inh

Slow inhibition Kinetics like GABA B

Slow inhibition Kinetics like GABA B G-protein concentration:

Slow inhibition Kinetics like GABA B G-protein concentration: Activated receptor concentration

Slow inhibition Kinetics like GABA B G-protein concentration: Activated receptor concentration Fast and slow Components:

Inhibitory neurons

Active currents:

Inhibitory neurons Active currents: Ca

Inhibitory neurons Active currents: Ca ( -> Ca spikes)

Inhibitory neurons Active currents: Ca K ( -> Ca spikes)

Inhibitory neurons Active currents: Ca K Ca-dependent K current ( -> Ca spikes)

Inhibitory neurons Active currents: Ca K Ca-dependent K current ( -> spike rate adaptation)( -> Ca spikes)

Inhibitory neurons Active currents: Ca K Ca-dependent K current Dynamics of n K(Ca) : ( -> spike rate adaptation)( -> Ca spikes)

Inhibitory neurons Active currents: Ca K Ca-dependent K current Dynamics of n K(Ca) : Ca dynamics: ( -> spike rate adaptation)( -> Ca spikes)

2 neurons (1 PN, 1LN)

6 PNs + 2 LNs

(fast) inhibition between LNs

6 PNs + 2 LNs (fast) inhibition between LNs

6 PNs + 2 LNs (fast) inhibition between LNs LNs take turns:

Full network (90+30)

Responses of 4 PNs to 1 stimulus

Reliable (trial-to-trial reproducible) firing timing when there is large Inhibitory input

Another stimulus: Input to same set of PNs but different LNs

Another stimulus: Input to same set of PNs but different LNs

Another stimulus: Input to same set of PNs but different LNs Same overall firing rate pattern, but different temporal fine structure

3 rd stimulus: Input to 90%-different set of neurons:

3 rd stimulus: Input to 90%-different set of neurons:

3 rd stimulus: Input to 90%-different set of neurons: Different firing pattern across neurons (but same network-average rate)

Blocking LN-LN inhibition LNs now spike ~ regularly

Blocking LN-LN inhibition LNs now spike ~ regularly Less difference between responses to stimuli 1 and 2

Reducing I K(Ca) (reducing LN spike-rate adaptation)

Reducing I K(Ca) (reducing LN spike-rate adaptation)

Reducing I K(Ca) Less precise timing, weaker temporal modulation, reduced discriminability (reducing LN spike-rate adaptation)

Role of slow LN-PN inhibition

Slow rate modulations abolished

Role of slow LN-PN inhibition Slow rate modulations abolished  reduced discriminability