Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity...

26
Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Spike-Timing-Dependent-Plasticity (STDP) models or how to understand memory. Pascal Helson INRIA Sophia-Antipolis - Advisor - Etienne Tanré Romain Veltz 15 mai 2017 Pascal Helson Spike-Timing-Dependent-Plasticity (STDP) models or how 1

Transcript of Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity...

Page 1: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Spike-Timing-Dependent-Plasticity (STDP) modelsor how to understand memory.

Pascal Helson

INRIA Sophia-Antipolis

- Advisor -Etienne TanréRomain Veltz

15 mai 2017

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.1

Page 2: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Table of content

1 Introduction to plasticity

2 Phenomenological STDP models

3 Our model

4 Simulations and perspectives

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.2

Page 3: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Neuron network functioning

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.3

Page 4: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Biophysical ModelsBasic cellular processes :

Pathways mediating functional synapticplasticity :

Too complex⇒Phenomenological models

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.4

Page 5: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Main idea of STDP

Hebb’s law (1949) :

”Neurons that fire together , wire together .”[?]

Spiking-Time Dependent Plasticity (STDP)[Bi and Poo, 1998]

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.5

Page 6: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Table of content

1 Introduction to plasticity

2 Phenomenological STDP models

3 Our model

4 Simulations and perspectives

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.6

Page 7: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Phenomenological STDP models

Simplify biophysical models

Neuron model

Plasticity model

Neuron network

Simulations

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.7

Page 8: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Plasticity rule

STDP learning rule

∆w = F+(w)G (∆t+)− F−(w)G (∆t−)

STDP experimental curve :

[Bi and Poo, 1998]

STDP deterministic curve (G) :

[Izhikevich and Desai, 2003]

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.8

Page 9: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Simulations

[Clopath et al., 2009]

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.9

Page 10: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Table of content

1 Introduction to plasticity

2 Phenomenological STDP models

3 Our model

4 Simulations and perspectives

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.10

Page 11: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Objectives

Rich enough to reproduce biological phenomena

Simple enough to be analysed mathematically

Take into account time scales differences

Adapted to simulation of not too small networks (10 000neurons)

Observe global properties of the network

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.11

Page 12: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Individual neuron model

Neurons are either at rest(0) orspiking(1).

Wt ∈ RN × RN synaptic weightsmatrix.

Vt ∈ {0, 1}N neuron system state.

Inhomogeneous jump rates :

0α′i (Wt ,Vt )

β

1

S : R 7→ R+∗ bounded, positive and

nondecreasing, αm > 0 :

α′i (Wt ,Vt) = S

(N∑i=1

W ijt V j

t

)+ αm

System with 2 neurons

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.12

Page 13: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Plasticity rule

Jump of ∆w = cste

If the neuron i spikes at t−, ∀j 6= i :

if s jt < δ then wij + ∆w with probability p+(s jt ) = 1s jt<δA+e− s

jt

τ+ ;

if s jt < δ then wji −∆w with probability p−(s jt ) = 1s jt<δA−e− s

jt

τ− .

STDP curve :

[Izhikevich and Desai, 2003]

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.13

Page 14: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Markov Process (Wt , St ,Vt)t>0 with (W0, S0,V0) = (w0, s0, v0) and :

Wt ∈ RN × RN synaptic weight matrix ;

St = (S1t , ..., S

Nt ) ∈ RN inter-arrival time between spikes ;

Vt ∈ {0, 1}N neuron system state.

State space : E = (RN × RN)× RN × {0, 1}N

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.14

Page 15: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Gi =

w + ∆w

0 . . . 0...

0 . . . 0~γ i

0 . . . 0...

0 . . . 0

︸ ︷︷ ︸

N×N matrix

+

0 . . . 0 0 . . . 0...

......

...

...... ~ζ i

......

......

......

0 . . . 0 0 . . . 0

, (~γ i , ~ζ i ) ∈ F i

F i =

(~γ i , ~ζ i ), ~γ i = (γ i1, ..., γ

iN), ~ζ i =

ζi1...ζ iN

, γ ij , ζ

ij ∈ {0, 1}, γ i

i = ζ ii = 0

For (~γ i , ~ζ i ) 6= (0, 0) :

φ(s, ~γ i , ~ζ i ) =∏j 6=i

[γ ij p

+(sj) + (1− γ ij )(1− p+(sj))

] [(ζ ij p

−(sj) + (1− ζ ij )(1− p−(sj))]

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.15

Page 16: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Generator of the process (Wt , St ,Vt)t>0

∼Cf (w , s, v) =

∑i

δ1(v i )β[f (w , s, v − ei )− f (w , s, v)]︸ ︷︷ ︸B↓f (w,s,v)

+ φ(s,w ,w)∑i

αi (w , v)δ0(v i ) (f (w , s − siei , v + ei )− f (w , s, v))︸ ︷︷ ︸B↑f (w,s,v)

+N∑i=1

∂si f (w , s, v)︸ ︷︷ ︸Btr f (w,s,v)

+∑i

αi (w , v)δ0(v i )

∑∼w∈Gi ,

∼w 6=w

(f (∼w , s − siei , v + ei )− f (w , s, v))φ(s,

∼w ,w)

︸ ︷︷ ︸

∼Af (w,s,v)

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.16

Page 17: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Assumption on time scales∑(~γ i ,~ζ i )6=(0,0) φ(s, ~γ i , ~ζ i )� φ(s, 0, 0)

⇒ (St ,Vt)t>0 change fast compare to (Wt)t>0

First results

Wt fixed, (St ,Vt)t>0 has a unique invariant measure πWt

One can compute the Laplace transform of πWt

Slow fast analysis gives the limit model for the weights :

Cf (w) =

∫R2+×{0,1}N

Af (w , s, v)πw (ds, dv)

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.17

Page 18: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Table of content

1 Introduction to plasticity

2 Phenomenological STDP models

3 Our model

4 Simulations and perspectives

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.18

Page 19: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Simulations2 neurons

Analytic

Numerical

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.19

Page 20: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Simulations2 neurons

Discontinuity of the density

��� ��� ���

�������������

������������� �������������

���

���

���

���

����

����

����

���

����

����

����

����

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.20

Page 21: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

SimulationsN neurons

STDP curve

-100 0 100-0.5

0.0

0.5

1.0

dt (ms)

dw/w

y1y2

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.21

Page 22: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Simulations2 neurons

Trajectories

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.22

Page 23: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Simulations10 neurons

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.23

Page 24: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Perspectives

Study the weights dynamics

Simulations to test with other plasticity rules

Neurons states from discrete to continuous.

Mean field approximations.

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.24

Page 25: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Bi, G.-q. and Poo, M.-m. (1998). Synaptic modifications in culturedhippocampal neurons : dependence on spike timing, synaptic strength, andpostsynaptic cell type. Journal of neuroscience, 18(24) :10464–10472.

Clopath, C., Büsing, L., Vasilaki, E., and Gerstner, W. (2009). Connectivityreflects coding : A model of voltage-based spike-timing-dependent-plasticitywith homeostasis. Nature.

Izhikevich, E. M. and Desai, N. S. (2003). Relating stdp to bcm. Neuralcomputation, 15(7) :1511–1523.

Morrison, A., Diesmann, M., and Gerstner, W. (2008). Phenomenologicalmodels of synaptic plasticity based on spike timing. Biological Cybernetics,98(6) :459–478.

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.25

Page 26: Spike-Timing-Dependent-Plasticity (STDP) models or how to ......Introduction to plasticity Phenomenological STDP models Our model Simulations and perspectives Table of content 1 Introductiontoplasticity

Introduction to plasticityPhenomenological STDP models

Our modelSimulations and perspectives

Thank you for your attentionDo you have questions ?

Pascal HelsonSpike-Timing-Dependent-Plasticity (STDP) models or how to understand memory.26