Reduced Neuron Models

Richard Naud, uOttawa

 Dendrites

Neuronal Excitability

Steps of information processing:

- Synapses: connection between neurons
- Dendrites: receive inputs
- Cell body: sums currents from dendrites
- Axon: sends to action potentials Cell body

How are action potential generated

given current flowing into cell body
from dendrites and synapses? Axon

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 2

 Biophysical description
I(t)
I(t) Current impinging on excitable
Ion channel
membrane patch Lipid
V(t) Membrane potential bilayer V(t)

I(t)
C Capacitance of the membrane
gL Conductance of the membrane V(t)
EL Equilibrium potential of ‘leak’

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 3

 Biophysical description
I(t)
I(t) Current impinging on excitable
Ion channel
membrane patch Lipid
V(t) Membrane potential bilayer V(t)

INa(t) IK(t) IH(t) IAHP(t)

Cm Capacitance of the membrane

dV
gL Conductance of the membrane I = C + gL(V − EL) + INa + IK + IH + IAHP
dt
EL Equilibrium potential of ‘leak’

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 4

 Leaky Integrate-and-Fire

INa(t) IK(t) IH(t) IAHP(t)

dV
I = Cm + gL(V − EL) + INa + IK + IH + IAHP
dt

Replace by a threshold for spike emission Ignore for now:

Followed by a reset to a fixed potential Action of ion channels other than
stereotypical action potential generation

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 5

 Leaky Integrate-and-Fire (LIF)
dV
Cm = − gL(V − EL) + I
dt Current
If V(t) = Vth then V(t + Δ) = EL I(t)

Membrane Potential
Subthreshold current step: Exponential
V(t)
relaxation to a steady-state.
I
V(t) = ( + EL)[1 − e −t/τm]
gL
τm = Cm /gL Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 6
 Leaky Integrate-and-Fire (LIF)
dV
Cm = − gL(V − EL) + I
dt Current
If V = Vth then V(t + Δ) = EL I(t)

V(t)
Suprathreshold current step:
Vth

Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 7
 Leaky Integrate-and-Fire (LIF)
dV
Cm = − gL(V − EL) + I
dt Current
If V = Vth then V(t + Δ) = EL I(t)
Spike ISI = interspike
V(t) ISI
interval (inverse
Suprathreshold current step: frequency)
Regular firing Vth

Δ Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 8
 Do Spikes Always Have the Same Shape?

Voltage reset
Spiking threshold
Variability across >100 spikes
Yes. Spikes follow stereotypical time course within 1-2 ms of onset.
Notable exception: spikes late in a high-frenquency burst.
Figure modified from: Mensi et al. J. Neurophys. (2012)
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 9
 Generalized Integrate-and-Fire

INa(t) IK(t) IH(t) IAHP(t)

dV
I=C + gL(V − EL) + INa + IK + IH + IAHP
dt

Replace by a threshold for spike emission Ignore for now:

Followed by a reset to a fixed potential Approximate with linearother
Action of ion channels dynamics
than
stereotypical action potential generation

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 10

 Generalized Integrate-and-Fire (GIF)

dV dwk Linearized
∑ k
Cm = − gL(V − EL) − w +I τk = ak(V − EL) − wk
dt k
dt subthreshold current
If V = Vth then V(t + Δ) = EL

ak : sub-threshold coupling of k’th current.

τk: time scale of k’th current
Linear dynamics

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 11

 Generalized Integrate-and-Fire (GIF)
Current
I(t)
Linearized sub-threshold currents
V(t) can capture some non-mono-
exponential features

Not a mono-exponential
relaxation

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 12

 Generalized Integrate-and-Fire (GIF)
Spike-frequency adaptation
Current Almost all cells show some spike-
I(t) frequency adaptation

Adaptation Modeled by adding a spike-

w(t) triggered inhibitory current with
linear dynamics
V(t) ISI

VT

Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 13
 Generalized Integrate-and-Fire (GIF)

dV dwk
∑ k
Cm = − gL(V − EL) − w +I τk = ak(V − EL) − wk
dt k
dt
Linearized spike-
If V = Vth then V(t + Δ) = EL wk(t + Δ) = wk + bk triggered current
bk: spike-triggered jump in k’th current.

Also, the threshold is allowed to jump up after every spike and decay to a baseline.

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 14

 Accuracy of Reduced Models

Real
To validate the neuron model we neuron

inject in-vivo-like current in neuron

and try to predict precise spike
timing (+/- 4 ms) using a
mathematical neuron model. Mathematical
model

Figure modified from: Gerstner, Kistler, Naud and Paninski Neuronal dynamics (2014)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 15

 Accuracy of Reduced Models
Model accuracy 100
[%]
75

Conclusion: Generalized 50
Integrate and Fire models 25
(GIF) are as accurate as
0
Hodgkin-Huxley (HH) LIF GIF HH GIF
models, despite being fast spiking
pyramidal cell interneuron
simpler

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 16

 Conclusion
Integrate-and-fire models form an 100
approximation to the complex 75
dynamics of real neurons.
50
Generalized Integrate-and-fire 25
models attain high accuracy by
0
taking into account adaptive features LIF GIF HH GIF
fast spiking
pyramidal cell interneuron

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 17

 Further information
Textbooks:
Gerstner, Kistler, Naud and Paninski, Neuronal Dyanmics (2014)
Tuckwell, Introduction to theoretical neurobiology vol 1 & 2 (1988)
Dayan and Abbott, Theoretical Neuroscience (2001)

GIF and extensions:

Richardson, Brunel, Hakim Journal of Neurophysiology (2003)
Fourcaud-Trocme et al. Journal of Neuroscience (2003)
Mensi et al. Journal of Neuorphysiology (2011)
Pozzorini et al. Nature Neuroscience (2013)
Mensi et al. PLoS Computational Biology (2016)

GIF accuracy:
Pillow et al. Journal of Neuroscience, (2005)
Pillow et al. Nature (2008)
Jolivet et al. J Neuroscience Methods (2009)
Naud and Gerstner, Science (2011)
Teeter et al. Nature Communications (2018)
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 1 18
The Statistics of Neuronal Responses
Richard Naud, uOttawa
 Characterization of the Input-Output Function

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 2

 Characterization of the Input-Output Function

Input Output

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 3

 Leaky Integrate and Fire - Constant Input

Firing Frequency as inverse

1
Current I interspike interval:
ISI

dV
C = − gL(V − EL) + I
Input dt Output

If V = θ then V(t + Δ) = Vr

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 4

 Leaky Integrate and Fire - Constant Input

Firing
frequency
1
ISI
Missing:
Formula

Current I

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 5

 Leaky Integrate and Fire - Noisy Input
Current I Firing Frequency as inverse of
1
average interspike interval:
⟨ISI⟩
+
Noise ξ

dV
C = − gL(V − EL) + I + ξ
Input dt Output

If V = θ then V(t + Δ) = Vr

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 6

 Leaky Integrate and Fire - Noisy Input

Firing
frequency
1
⟨ISI⟩
Missing:
Formula

Current I

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 7

 Leaky Integrate and Fire - Noisy Input
Current I
σT
CV
+ ⟨ISI⟩
Noise ξ

dV
C = − gL(V − EL) + I + ξ
Input dt Output

If V = θ then V(t + Δ) = Vr

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 8

 Leaky Integrate and Fire - Noisy Input
σT
CV
⟨ISI⟩

Missing:
Formula

Current I

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 9

 Correlation of Pairs

Correlation
Current I Noise 1 ξ ρ MISSING EQUATION
LIF
Noise 2 ξ

Current I Noise 3 ξ LIF

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 10

 Correlation of Pairs
Correlation
ρ

Missing:
Formula

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 11

 Ensemble Response
Current I Noise 1 ξ LIF

PSTH
LIF ρ MISSING EQUATION
…
… LIF

Current I Noise 3 ξ LIF

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 12

 Leaky Integrate and Fire - Noisy Input

PSTH ν(t)

Time t Current I

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 13

 Transmission of Output Statistics
LIF

LIF
LIF

LIF

LIF

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 14

 Transmission of Output Statistics

What comes next requires synapse

models

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 15

 Conclusion

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 16

Synaptic Transmission
Richard Naud, uOttawa
 Transmission

How does a the firing of one

given neuron affect the neuron
connected to it?

What statistical property is

transmitted?

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3 2

 Synaptic Currents

Pre-synaptic: Action
Vesicle
potential arrives at the release
synapse and triggers
transmitter release
Post-synaptic: transmitter
binds to receptor and Transmitter-
gated ion
triggers changes in ionic
channel
conductance

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Glutamatergic Gabaergic
Synaptic Currents
A presynaptic spike can cause
both fast and slow post-
synaptic currents (PSCs)

Glutamatergic synapses: release Gabaergic synapses: release

glutamate, causing excitatory GABA, causing inhibitory PSCs
PSCs (EPSCs). (IPSCs)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Synaptic Currents
Currents are mediated by
conductance changes:

IAMPA(t) = gAMPA(t)(V − EE ) gAMPA ∼ Θ(t − tf )e −(t−tf )/τAMPA

INMDA(t) = gNMDA(V, t)(V − EE ) All others: g(t) is a bi-

exponential function

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Current vs Conductance vs Potential
IAMPA(t) = gAMPA(t)(V − EE ) At resting At reversal
potential potential

Excitatory post-synaptic conductance (EPSC)

Excitatory post-synaptic current (EPSC)

Shunting due to
Smoothing
Excitatory post-synaptic potential (EPSP) membrane time constant

Because reversal potential is typically well above firing threshold: excitatory

conductance typically proportional to current IAMPA ≈ αgAMPA
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Synaptic bombardment
Neuron 1

Neuron 2

Neuron 3

Post-synaptic neuron

Post-synaptic potential

V(t)
== = ϵ1ϵ(t1(t
ϵ−i(t−
tf −
)tf )tf )
∑∑∑
V(t)
V(t)
f f i
Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Ensemble Rate - Peri-Stimulus Time-Histogram
PSTH
1 1
ϵ(t − tf(i), Δt)
N Δt ∑
ν(t) =
i

Raster plot: each line is PSTH: An average across

neuron, each dot is a spike columns of the raster plot

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2

 Ensemble Rate - Peri-Stimulus Time-Histogram
Ensemble rate
1 1
ϵ(t − tf(i), Δt)
N Δt ∑
ν(t) =
i
Equivalent!
Synapses communicate the
ensemble rate of the pre-
synaptic population

∑∑
V(t) = ϵi(t − tf )
f i
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2
 Can Spikes be Shuffled across Population?
Neuron 1

Neuron 2

Neuron 3

Post-synaptic neuron

Post-synaptic potential

∑∑
V(t) = ϵi(t − tf )
NO CHANGE! ?
f i
Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Short-Term Dynamics

The amplitude of individual post-synaptic Facilitation

conductance depends on the history of
firing on fast (0.1-5 s) timescale

Some pathways make strictly facilitating synapses, Depression

others strictly depressing synapses

These distinct dynamics can be generated from the

same axon

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Distinct types of synapses

Facilitating: Firing causes Depressing: Firing causes a

increase in vesicle release decrease in size of readily
probability releasable pool

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Tsodyks Markram Model

PSC amplitude is conceived as a multiplication of two factors: uR

dR 1 − R Depressing: Firing causes a decrease

= if spike then R ← uR in R , which will recover to 1 on a time
dt τR
scale τR
Facilitating: Firing causes increase in
du U − u
= if spike then u ← f(1 − u) u controlled by by jump f and
dt τu
timescale τu , but ensuring it remains
between 0 and 1
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Tsodyks Markram Model

Example: fast τR , slower τuand high f leads to facilitation

Saturation of
jumps uR but jumps as u
decays fast increases
creates a
saturation of
jumps f (1 − u) facilitation

Figures: Rossbroich Trotter et al.. BioRxiv (2020)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Can Spikes be Shuffled across Population?

Neuron 1 Facilitating synapses

Neuron 2

Neuron 3

Not Equivalent!
Dynamic synapses do
Post-synaptic potential
not communicate
ensemble rate
Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3
 Conclusion

Static synapse: cummunicate

ensemble rate of pre-synaptic
neurons

Dynamic synapse: introduce

sensitivity to single-neuron
interspike intervals

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 3

 Further Information
Textbooks:
Gerstner, Kistler, Naud and Paninski, Neuronal Dyanmics (2014)
Koch, Biophysics of Computation (2004)
Dayan and Abbott, Theoretical Neuroscience (2001)

Short-term dynamics
Tsodyks and Markram PNAS (1997)
Markram and Tsodyks PNAS (1998)
Hennig Frontiers in Computational Neuroscience (2013)
Naud and Sprekeler PNAS (2018)
Rossbroich, Trotter et al. BioRxiv (2020)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 17

Spike Timing and Plasticity
Richard Naud, uOttawa
 Plasticity
The connection strength of a
synapse is plastic, malleable. It is
is sculpted by patterns of activity.

Synaptic plasticity if often loosely

described by ‘what fires together
wires together’, but does this
mean that ‘what spikes together
wires together’?

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 2

 Synaptic Weights
The connection strength, W, is pre
the amplitude of the PSC. ΔW
Long-Term Potentiation (LTP):
W
plasticity-inducing protocol,
causing the connection strength post
to increase ΔW > 0 .
Long-Term Depression (LTD):
The connection strength is
decreased ΔW < 0.

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 3

 Plasticity Factors
Relative timing Neuromodulation Firing pattern

ΔW
ΔW ΔW

Relative
Concentration Burstiness
Δt

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 4

 Modelling Spike-Timing-Dependent Plasticity
Experimental Data

pre
In-vivo like spike trains

post

ΔW How should W evolve?

Nearest neighbour?
Triplets? How?
Δt
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 5
 Modelling Spike-Timing-Dependent Plasticity
Pre- and post-synaptic traces
pre Pre-synaptic trace is thought to be the activation of a
metabotropic receptor, or a pathway triggered by spike
voltage: triggered by pre-synaptic spike and evanescent
Pi(t) A+
post Post-synaptic trace is thought to be the voltage trace from
the back-propagating action potential to the location of the
synapse: triggered by post-synaptic spike and evanescent
M(t) A−

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 6

 Modelling Spike-Timing-Dependent Plasticity
Pre- and post-synaptic traces
pre At each pre-synaptic spike, change
weight according post-synaptic trace

Pi(t) ΔW(tpre) = M(tpre)W(tpre)

post

M(t)

W(t)
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 7
 Modelling Spike-Timing-Dependent Plasticity
Pre- and post-synaptic traces
pre At each pre-synaptic spike, change
weight according post-synaptic trace

Pi(t) ΔW(tpre) = M(tpre)W(tpre)

post At each post-synaptic spike, change

weight according post-synaptic trace

M(t) ΔW(tpost) = Pi(tpost)W(tpost)

W(t) LTD LTP

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 8
 Modelling Spike-Timing-Dependent Plasticity
Pre- and post-synaptic traces Controlled by
τtrace = 20 ms
pre
ΔW Controlled by
Pi(t)
A+
post Controlled by
A− Δt
M(t)

W(t) LTD LTP

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 9
 Modelling Spike-Timing-Dependent Plasticity
Synaptic bounds
pre At each pre-synaptic spike, change weight according
post-synaptic trace or up to upper bound

Pi(t) ΔW(t
ΔW(t
pre)pre
=) min(M(t
= M(tprepre
)W(t
)W(t ) ),1)
prepre

post At each post-synaptic spike, change weight according post-

synaptic trace or down to zero

M(t) ΔW(tpost) = max(Pi(tpost)W(tpost),0)

W(t)
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 10
 Exploring Spike-Timing-Dependent Plasticity
Start simulation with weak connections and silent post 1000 excitatory 200 inhibitory
synaptic neuron
No pre-post

{
Coincidence

pre …
Synapses 1 postsynaptic
No plasticity with STDP neuron

post
Neuron remains
silent
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 11
 Exploring Spike-Timing-Dependent Plasticity
Start simulation with stronger connections and active 1000 excitatory 200 inhibitory
post synaptic neuron
Regular firing

pre
{ … ΔW
Random pre-post timing

Δt
1 postsynaptic
neuron

post Net depression

if A−>A+

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 12

 Exploring Spike-Timing-Dependent Plasticity
Net potential
Current
Net excitatory
1000 excitatory 200 inhibitory
‘Balance of excitation
and inhibition’ Firing
Threshold

0
‘Fluctuation-driven’
Regime 1 postsynaptic
neuron

Net inhibitory

Time
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 13
 Exploring Spike-Timing-Dependent Plasticity
Start simulation with stronger connections and active 1000 excitatory 200 inhibitory
post synaptic neuron Regular firing

pre
{ … Random pre-post timing

Net depression
1 postsynaptic
neuron

post

Irregular firing
Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 14
 Exploring Spike-Timing-Dependent Plasticity
STDP can stabilize net excitation at the fluctuation-
1000 excitatory 200 inhibitory
driven regime
post CV I/E
slope of 1
rate
[Hz] 20 1 1

10 0.5 0.5
1 postsynaptic
neuron

10 40 10 40 10 40

pre rate [Hz] pre rate [Hz] pre rate [Hz]

Redrawn from : Song, Miller and Abbott Nature Neuroscience (2000)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 4 15
 Further Information
Textbooks:
Gerstner, Kistler, Naud and Paninski, Neuronal Dynamics (2014)
Dayan and Abbott, Theoretical Neuroscience (2001)

Spike-Timing Dependent Plasticity (STDP)

Gerstner et al. Nature (1996)
Kempter et al. Phys Rev E (1999)
Song, Miller and Abbott Nature Neuroscience (2000)
Song and Abbott Neuron (2001)
Vogels et al. Science (2011)

Neuromodulation and Firing Patterns

Izhikevich Celrebral Cortex (2007)
Payeur et al. BioRxiv (2020)
Pawlak et al. Front. Synap. Neurosci. (2010)

Richard Naud ⦁ Neuron and Synapse Models Week 3 ⦁ Day 1 ⦁ Tutorial 2 16