Part II: Population Models
description
Transcript of Part II: Population Models
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Part II: Population Models
BOOK: Spiking Neuron Models, W. Gerstner and W. Kistler Cambridge University Press, 2002
Chapters 6-9
Laboratory of Computational Neuroscience, LCN, CH 1015 LausanneSwiss Federal Institute of Technology Lausanne, EPFL
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Chapter 6: Population Equations
BOOK: Spiking Neuron Models, W. Gerstner and W. Kistler Cambridge University Press, 2002
Chapter 6
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10 000 neurons3 km wires
1mm
Signal:action potential (spike)
action potential
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Spike Response Model
iuij
fjtt
Spike reception: EPSP
fjtt
Spike reception: EPSP
^itt
^itt
Spike emission: AP
fjtt ^
itt tui j f
ijw
tui Firing: tti ^
linear
threshold
Spike emission
Last spike of i All spikes, all neurons
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Integrate-and-fire Model
iui
fjtt
Spike reception: EPSP
)(tRIuudtd
ii
tui Fire+reset
linear
threshold
Spike emission
resetI
j
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escape process (fast noise)
parameter changes (slow noise)
stochastic spike arrival (diffusive noise)
Noise models
A B C
u(t)
noise
white(fast noise)
synapse(slow noise)
(Brunel et al., 2001)
t
t
dttt^
)')'(exp()( )¦( ^ttPI
: first passagetime problem
)¦( ^ttPI Interval distribution
^t ^t ^tt
Survivor functionescape rate
)(t
))(()( tuftescape rate stochastic reset
)¦( ^ttPI )( fttG
Interval distribution
Gaussian about ft
)(tRIudtdu
ii
noisy integration
ft
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Homogeneous Population
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populations of spiking neurons
I(t)
?
population dynamics? t
t
tNtttntA
);()(population
activity
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Homogenous network (SRM)
NJwij
0
Spike reception: EPSP
Spike emission: AP
fjtt ^
itt tui j f
ijwLast spike of i All spikes, all neurons
fjtt
^itt
Synaptic coupling
potential
fullyconnected N >> 1
dsstIs )( external input
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NJwij
0
fjtt ^
itt tui j f
ijwLast spike of i All spikes, all neuronspotential
dsstIs )( external input
dsstIs )( tui ^itt dsstAsJ )(0
potential
^tt ^| ttu )(thinput potential
fullyconnected
refractory potential
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Homogenous network
Response to current pulse
Spike emission: AP
s
^itt
potential
^tt ^| ttu )(thinput potential
itt ˆ tui
Last spike of ipotential
dsstIsJ )(0 external input
dsstAs )( Population activity
All neurons receive the same input
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Assumption of Stochastic spike arrival: network of exc. neurons, total spike arrival rate A(t)
)()(,
0 fk
fkrest tt
Cq
NJ
uuudtd
u
0u
EPSC
Synaptic current pulses
Homogeneous network (I&F)
)()( tIRuuudtd
rest
)()( 0 tAqJtI
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Density equations
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Stochastic spike arrival: network of exc. neurons, total spike arrival rate A(t)
)()()(,
0 f
feext
fk
fk
erest tt
Cq
JttCq
NJ
uuudtd
u
0u
EPSC
Synaptic current pulses
Density equation (stochastic spike arrival)
)()()( ttIRuuudtd
rest Langenvin equation,Ornstein Uhlenbeck process
fqJtAqJtI ext )()( 0
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u
p(u)
Density equation (stochastic spike arrival)
u
Membrane potential density
)()(),()],()([),( 2
22
21 tAuutup
utupuV
utup
t r
Fokker-Planck
drift diffusionAqJuuV 0)(
kkk w22
spike arrival rate
source term at reset
A(t)=flux across threshold
utupu
tA ),()(
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Integral equations
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tdtAttPtA I
t
ˆ)ˆ(ˆ|)(
Population Dynamics
tdtAttSI
t
ˆ)ˆ(ˆ|1
Derived from normalization
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Escape Noise (noisy threshold)
t̂
)(t
I&F with reset, constant input, exponential escape rate
Interval distribution
t̂
)ˆ(0 ttP )')ˆ'(exp()ˆ()ˆ( ̂t
t
dtttttttP
)exp())ˆ(()ˆ()ˆ(u
ttuttuftt
escape rate
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tdtAttPtA I
t
ˆ)ˆ(ˆ|)(
Population Dynamics
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Wilson-Cowan
population equation
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escape process (fast noise)
Wilson-Cowan model
h(t)
^t t
)(t
))(()( thftescape rate
(i) noisy firing(ii) absolute refractory time
abs
))(()( thftA
population activity
t
t abs
dttA
]')'(1[
(iii) optional: temporal averaging
))(()()( thgtAtAdtd
abs
abs
ttfor
ttforthftuft
)ˆ(00
)ˆ())(())(()(
escape rate
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escape process (fast noise)
Wilson-Cowan model
h(t)
^t t
)(t
(i) noisy firing(ii) absolute refractory time
abs
))(()( thftA
population activity
t
t abs
dttA
]')'(1[
abs
abs
ttfor
ttforthftuft
)ˆ(00
)ˆ())(())(()(
escape rate
tdtAttPtA I
t
ˆ)ˆ(ˆ|)(
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Population activity in spiking neurons (an incomplete history)
1972 - Wilson&Cowan; Knight Amari
1992/93 - Abbott&vanVreeswijk Gerstner&vanHemmen
Treves et al.; Tsodyks et al. Bauer&Pawelzik
1997/98 - vanVreeswijk&Sompoolinsky Amit&Brunel Pham et al.; Senn et al.
1999/00 - Brunel&Hakim; Fusi&Mattia Nykamp&Tranchina Omurtag et al.
Fast transientsKnight (1972), Treves (1992,1997), Tsodyks&Sejnowski (1995)Gerstner (1998,2000), Brunel et al. (2001), Bethge et al. (2001)
Integral equation
Mean field equationsdensity (voltage, phase)
Heterogeneous netsstochastic connectivity
(Heterogeneous, non-spiking)
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Chapter 7: Signal Transmission and Neuronal Coding
BOOK: Spiking Neuron Models, W. Gerstner and W. Kistler Cambridge University Press, 2002
Chapter 7
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Coding Properties of Spiking Neuron ModelsCourse (Neural Networks and Biological Modeling) session 7 and 8
Laboratory of Computational Neuroscience, LCN, CH 1015 LausanneSwiss Federal Institute of Technology Lausanne, EPFL
PSTH(t)
500 trials
I(t)
forward correlationfluctuating input
I(t)reverse correlationProbability of
output spike ?
I(t) A(t)?0t
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Theoretical Approach
- population dynamics
- response to single input spike (forward correlation)
- reverse correlations
A(t)
500 neurons
PSTH(t)
500 trials
I(t) I(t)
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Population of neurons
h(t)
I(t) ?0t
))(()( thgtA
A(t)
A(t)
A(t)
))(()( tIgtA
))('),(()( tItIgtA
potential
A(t) ))(()()( thgtAtAdtd
t
tNtttntA
);()(population
activity
N neurons,- voltage threshold, (e.g. IF neurons)- same type (e.g., excitatory) ---> population response ?
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Coding Properties of Spiking Neurons:
Laboratory of Computational Neuroscience, LCN, CH 1015 LausanneSwiss Federal Institute of Technology Lausanne, EPFL
- forward correlations- reverse correlations
1. Transients in Population Dynamics - rapid transmission2. Coding Properties
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Example: noise-free
tdtAttPtA I
t
ˆ)ˆ(ˆ|)(
))ˆ(ˆ(ˆ| tTttttPI
)(tA )( TtA
''1
uh
Population Dynamics
I(t) h’>0h(t)
T(t^)
higher activity
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noise-free
Theory of transients
)(tA )( TtA
''1
uh
I(t)h(t)
I(t) ?0t
potential dsstIs )( ^tt ^| ttu
)(thinput potential 0)(' ttth
)()( 00 ttAAtA A(t)
External input.No lateral coupling
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Theory of transients A(t)
no noise
I(t)h(t)
noise-free
noise model B
slow noise
I(t)h(t)
(reset noise)
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u
p(u)
u
Membrane potential density
Hypothetical experiment: voltage step
u
p(u)
Immediate responseVanishes linearly
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Transients with noise
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escape process (fast noise)
parameter changes (slow noise)
stochastic spike arrival (diffusive noise)
Noise models
A B C
u(t)
noise
white(fast noise)
synapse(slow noise)
(Brunel et al., 2001)
t
t
dttt^
)')'(exp()(
)¦( ^ttPI Interval distribution
^t ^t ^tt
Survivor functionescape rate
)(t
))(()( tuftescape rate stochastic reset
)¦( ^ttPI )( fttG
Interval distribution
Gaussian about ft
)(tRIudtdu
ii
noisy integration
ft
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Transients with noise:Escape noise (noisy threshold)
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linearize
tdtAttPtA I
t
ˆ)ˆ(ˆ|)(
)()( 0 tAAtA
Theory with noise A(t)
)()( 0 thhth
I(t)h(t)
0A
dsstIsth )()()(
sA 1
0 inverse mean interval
I
Llow noiselow noise: transient prop to h’
high noise: transient prop to h
h: input potential
high noise
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Theory of transients A(t)
low noise
I(t)h(t)
noise-free
(escape noise/fast noise) noise model A
low noise
fast
noise model A
I(t)h(t)
(escape noise/fast noise)
high noise
slow
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Transients with noise:Diffusive noise (stochastic spike arrival)
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escape process (fast noise)
parameter changes (slow noise)
stochastic spike arrival (diffusive noise)
Noise models
A B C
u(t)
noise
white(fast noise)
synapse(slow noise)
(Brunel et al., 2001)
t
t
dttt^
)')'(exp()(
)¦( ^ttPI Interval distribution
^t ^t ^tt
Survivor functionescape rate
)(t
))(()( tuftescape rate stochastic reset
)¦( ^ttPI )( fttG
Interval distribution
Gaussian about ft
)(tRIudtdu
ii
noisy integration
ft
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u
p(u)
Diffusive noiseu
Membrane potential density
p(u)
Hypothetical experiment: voltage step
Immediate responsevanishes quadratically
),(
)],()([
),(
2
22
21 tup
u
tupuAu
tupt
Fokker-Planck
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u
p(u)
SLOW Diffusive noiseu
Membrane potential density
Hypothetical experiment: voltage step
Immediate responsevanishes linearly
p(u)
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Signal transmission in populations of neurons
Connections4000 external4000 within excitatory1000 within inhibitory
Population- 50 000 neurons- 20 percent inhibitory- randomly connected
-low rate-high rate
input
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Population- 50 000 neurons- 20 percent inhibitory- randomly connected
Signal transmission in populations of neurons
100 200time [ms]
Neuron # 32374
50
u [mV]
100
0
10A [Hz]
Neu
ron
#
32340
32440
100 200time [ms]50
-low rate-high rate
input
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Signal transmission - theory
- no noise
- slow noise (noise in parameters)
- strong stimulus
- fast noise (escape noise) prop. h(t) (potential)
prop. h’(t) (current)
See also: Knight (1972), Brunel et al. (2001)
fast
slow
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Transients with noise: relation to experiments
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Experiments to transients A(t)
V1 - transient response
V4 - transient response
Marsalek et al., 1997
delayed by 64 ms
delayed by 90 ms
V1 - single neuron PSTH
stimulus switched on
Experiments
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input A(t)
A(t)
A(t)
A(t)
See also: Diesmann et al.
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How fast is neuronal signal processing?
animal -- no animalSimon ThorpeNature, 1996
Visual processing Memory/association Output/movement
eye
Reaction time experiment
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How fast is neuronal signal processing?
animal -- no animalSimon ThorpeNature, 1996
Reaction time
Reaction time
# ofimages
400 msVisual processing Memory/association Output/movement
Recognition time 150ms
eye
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Coding properties of spiking neurons
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Coding properties of spiking neurons
- response to single input spike
(forward correlations)
A(t)
500 neurons
PSTH(t)
500 trials
I(t)I(t)
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Coding properties of spiking neurons
- response to single input spike
(forward correlations)
I(t) Spike ?Two simple arguments1)
2)
Experiments: Fetz and Gustafsson, 1983 Poliakov et al. 1997
(Moore et al., 1970)
PSTH=EPSP
(Kirkwood and Sears, 1978)
PSTH=EPSP’
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Forward-Correlation Experiments A(t)
Poliakov et al., 1997
I(t) PSTH(t)
1000 repetitionsnoise
high noise low noiseprop. EPSP prop. EPSP
ddt
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^^^ )(|)( dttAttPtA I
t
Population Dynamics
)()( 0 thhth h: input potential dsstIsth )()()(
A(t) PSTH(t)I(t)I(t)
full theory
linear theory
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Forward-Correlation Experiments A(t)
Theory: Herrmann and Gerstner, 2001
high noise low noisePoliakov et al., 1997
high noise low noise
blue: full theoryred: linearized theory
blue: full theoryred: linearized theory
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Forward-Correlation Experiments A(t)
Poliakov et al., 1997
I(t) PSTH(t)
1000 repetitionsnoise
high noise low noiseprop. EPSP prop. EPSP
ddt
prop. EPSP
prop. EPSPddt
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Reverse Correlations
Laboratory of Computational Neuroscience, LCN, CH 1015 LausanneSwiss Federal Institute of Technology Lausanne, EPFL
fluctuating input
I(t)
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Reverse-Correlation Experiments
after 1000 spikes
)(tI
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)()( 0 thhth h: input potential dsstIsth )()()(
Linear Theory
Fourier Transform
)(~)(~)(~ IGA
0
)()()( dsstIsGtA
Inverse Fourier Transform
)(~1)(~)(~
)(~ 0
PLAiG
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Signal transmissionI(t) A(t)
)()()(
fIfAfG T=1/f
(escape noise/fast noise) noise model A
low noise
high noise
noise model B (reset noise/slow noise)
high noiseno cut-off
low noise
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Reverse-Correlation Experiments (simulations)
after 1000 spikes
0
)()()( dsstIsGtA
theory:G(-s)
)(tI
after 25000 spikes
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Laboratory of Computational Neuroscience, EPFL, CH 1015 Lausanne
Coding Properties of spiking neurons
I(t)
?
- spike dynamics -> population dynamics- noise is important - fast neurons for slow noise - slow neurons for fast noise
- implications for - role of spontaneous activity - rapid signal transmission - neural coding - Hebbian learning
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Chapter 8: Oscillations and Synchrony
BOOK: Spiking Neuron Models, W. Gerstner and W. Kistler Cambridge University Press, 2002
Chapter 8
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Stability of Asynchronous State
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Stability of Asynchronous State
Search for bifurcation points
linearize
^^^ )(|)( dttAttPtA I
t
)()( 0 tAAtA )()( 0 thhth dsstAsJth )()()(
h: input potential
A(t)
ttieAtA 1)(0
fully connected coupling J/N
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Stability of Asynchronous State A(t)
delayperiod
)()( sess0 for
stable0
03
02
noise
T
)(s
s
)(s
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Stability of Asynchronous State s
)(s
ms0.1
ms2.1
4.1
ms0.2
ms0.3
ms4.0
06
05
04
03
02
T 2
0
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Chapter 9: Spatially structured networks
BOOK: Spiking Neuron Models, W. Gerstner and W. Kistler Cambridge University Press, 2002
Chapter 9
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Continuous Networks
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)(tAi
Several populations
i k
Continuum
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)(),( AtA
Continuum: stationary profile