convergence and stability in networks with spiking neurons stan gielen dept. of biophysics magteld...
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Convergence and stability in networks with spiking neurons
Stan Gielen
Dept. of BiophysicsMagteld Zeitler
Daniele Marinazzo
V mV
0 mV
V mV
0 mV
IC
INa
Membrane voltage equation
-Cm dV/dt = gmax, Nam3h(V-Vna) + gmax, K n4 (V-VK ) + g leak(V-Vleak)
K
V (mV)
mmOpen Closedm
m
mProbability:
State:
(1-m)
Channel Open Probability:
dt
dm m)1( m m m
hhdtdh
hh )1(
Gating kinetics
m.m.m.h=m3h
mm
mm
mm
1
Simplification of Hodgkin-HuxleyFast variables• membrane potential V• activation rate for Na+
m
Slow variables• activation rate for K+ n• inactivation rate for
Na+ h
-C dV/dt = gNam3h(V-Ena)+gKn4(V-EK)+gL(V-EL) + I
dm/dt = αm(1-m)-βmm
dh/dt = αh(1-h)-βhh
dn/dt = αn(1-n)-βnn
Morris-Lecar model
Phase diagram for the Morris-Lecar model
Linearisation around singular point :
W
V
b
V
W
V
dt
d
1)1( 2
*
*
WWW
VVV
Overview• What’s the fun about synchronization ?• Neuron models• Phase resetting by external input• Synchronization of two neural oscillators• What happens when multiple oscillators are coupled ?• Feedback between clusters of neurons• Stable propagation of synchronized spiking in neural
networks• Current problems
Neuronal synchronizationT
ΔTΔ(θ)= ΔT/T
Phase shift as a function of the relative phase of the external input.
Phase advance
Hyperpolarizing stimulus
Depolarizing stimulus
Neuronal synchronizationT
ΔTΔ(θ)= ΔT/T
Suppose:
• T = 95 ms
• external trigger: every 76 ms
• Synchronization when ΔT/T=(95-76)/95=0.2
• external trigger at time 0.7x95 ms = 66.5 ms
ExampleT=95 ms
P=76 ms = T(95 ms) - Δ(θ)
For strong excitatory coupling, 1:1 synchronization is not unusual. For weaker coupling we may find other rhythms, like 1:2, 2:3, etc.
Neuronal synchronizationT
ΔTΔ(θ)= ΔT/T
Suppose:
• T = 95 ms
• external trigger: every 76 ms
• Synchronization when ΔT/T=(95-76)/95=0.2
• external trigger at time 0.7x95 ms = 66.5 ms
StableUnstable
Convergence to a fixed-point Θ* requires
Substitution of and expansion near gives
Convergence requires
and constraint gives
TPnnn /)(1
TPnnn /)(1
TP /)( * |||| **
1 nn
n= n* *
nn 1 nn TP
)(
/)(
)( *
1)(1
n
n n
n
1 <1
-1< 1)(<1 and so –2 < )(<0
T
P
Overview• What’s the fun about synchronization ?• Neuron models• Phase resetting by external input• Synchronization of two neural oscillators• What happens when multiple oscillators are coupled ?• Feedback between clusters of neurons• Stable propagation of synchronized spiking in neural
networks• Current problems