izhikevich_psc_alpha¶

izhikevich_psc_alpha - Detailed Izhikevich neuron model with alpha-kernel post-synaptic current

Description¶

Implementation of the simple spiking neuron model introduced by Izhikevich 1, with membrane potential in (milli)volt and current-based synapses.

The dynamics are given by:

\[ \begin{align}\begin{aligned}C_m \frac{dV_m}{dt} = k (V - V_t)(V - V_t) - u + I + I_{syn,ex} + I_{syn,in} \frac{dU_m}{dt} = a(b(V_m - E_L) - U_m)\\\begin{split}&\text{if}\;\;\; V_m \geq V_{th}:\\ &\;\;\;\; V_m \text{ is set to } c &\;\;\;\; U_m \text{ is incremented by } d\end{split}\end{aligned}\end{align} \]

On each spike arrival, the membrane potential is subject to an alpha-kernel current of the form:

\[I_syn = I_0 \cdot t \cdot \exp\left(-t/\tau_{syn}\right) / \tau_{syn}\]

References¶

1: Izhikevich, Simple Model of Spiking Neurons, IEEE Transactions on Neural Networks (2003) 14:1569-1572

Authors¶

Hanuschkin, Morrison, Kunkel

Parameters¶

Name	Physical unit	Default value	Description
C_m	pF	200.0pF	Membrane capacitance
k	pF / (ms mV)	8.0pF / mV / ms	Spiking slope
V_r	mV	-65.0mV	resting potential
V_t	mV	-45.0mV	threshold potential
a	1 / ms	0.01 / ms	describes time scale of recovery variable
b	nS	9.0nS	sensitivity of recovery variable
c	mV	-65mV	after-spike reset value of V_m
d	pA	60.0pA	after-spike reset value of U_m
V_peak	mV	0.0mV	Spike detection threashold (reset condition)
tau_syn_ex	ms	0.2ms	Synaptic Time Constant Excitatory Synapse
tau_syn_in	ms	2.0ms	Synaptic Time Constant for Inhibitory Synapse
t_ref	ms	2.0ms	Refractory period
I_e	pA	0pA	constant external input current

State variables¶

Name	Physical unit	Default value	Description
V_m	mV	-65mV	Membrane potential
U_m	pA	0pA	Membrane potential recovery variable

Equations¶

\[\frac{ dV_{m} } { dt }= \frac 1 { C_{m} } \left( { (k \cdot (V_{m} - V_{r}) \cdot (V_{m} - V_{t}) - U_{m} + I_{e} + I_{stim} + I_{syn,inh} + I_{syn,exc}) } \right)\]

\[\frac{ dU_{m} } { dt }= a \cdot (b \cdot (V_{m} - V_{r}) - U_{m})\]

Source code¶neuron izhikevich_psc_alpha:

  state:
    r integer = 0 # number of steps in the current refractory phase
    V_m mV = -65 mV # Membrane potential
    U_m pA = 0 pA   # Membrane potential recovery variable
  end

  equations:
    # synapses: alpha functions
    kernel I_syn_in = (e/tau_syn_in) * t * exp(-t/tau_syn_in)
    kernel I_syn_ex = (e/tau_syn_ex) * t * exp(-t/tau_syn_ex)

    inline I_syn_exc pA = convolve(I_syn_ex, spikesExc)
    inline I_syn_inh pA = convolve(I_syn_in, spikesInh)

    V_m' = ( k * (V_m - V_r) * (V_m - V_t) - U_m + I_e + I_stim + I_syn_inh + I_syn_exc ) / C_m
    U_m' = a * ( b*(V_m - V_r) - U_m )
  end

  parameters:
    C_m pF = 200. pF           # Membrane capacitance
    k pF/mV/ms = 8. pF/mV/ms   # Spiking slope
    V_r mV = -65. mV           # resting potential
    V_t mV = -45. mV           # threshold potential
    a 1/ms = 0.01 /ms          # describes time scale of recovery variable
    b nS = 9. nS               # sensitivity of recovery variable
    c mV = -65 mV              # after-spike reset value of V_m
    d pA = 60. pA              # after-spike reset value of U_m
    V_peak mV = 0. mV          # Spike detection threashold (reset condition)
    tau_syn_ex ms = 0.2 ms     # Synaptic Time Constant Excitatory Synapse
    tau_syn_in ms = 2.0 ms     # Synaptic Time Constant for Inhibitory Synapse
    t_ref ms = 2.0 ms          # Refractory period

    # constant external input current
    I_e pA = 0 pA
  end

  internals:
    RefractoryCounts integer = steps(t_ref) # refractory time in steps
  end

  input:
    spikesInh pA <- inhibitory spike
    spikesExc pA <- excitatory spike
    I_stim pA <- continuous
  end

  output: spike

  update:
    integrate_odes()

    # refractoriness and threshold crossing
    if r > 0: # is refractory?
      r -= 1
    elif V_m >= V_peak:
      V_m = c
      U_m += d
      emit_spike()
      r = RefractoryCounts
    end

  end

end