hh_cond_exp_traub

hh_cond_exp_traub - Hodgkin-Huxley model for Brette et al (2007) review

Description

hh_cond_exp_traub is an implementation of a modified Hodgkin-Huxley model.

This model was specifically developed for a major review of simulators 1, based on a model of hippocampal pyramidal cells by Traub and Miles 2. The key differences between the current model and the model in 2 are:

This model is a point neuron, not a compartmental model.
This model includes only I_Na and I_K, with simpler I_K dynamics than in 2, so it has only three instead of eight gating variables; in particular, all Ca dynamics have been removed.
Incoming spikes induce an instantaneous conductance change followed by exponential decay instead of activation over time.

This model is primarily provided as reference implementation for hh_coba example of the Brette et al (2007) review. Default parameter values are chosen to match those used with NEST 1.9.10 when preparing data for 1. Code for all simulators covered is available from ModelDB 3.

Note: In this model, a spike is emitted if \(V_m >= V_T + 30\) mV and \(V_m\) has fallen during the current time step.

To avoid that this leads to multiple spikes during the falling flank of a spike, it is essential to choose a sufficiently long refractory period. Traub and Miles used \(t_{ref} = 3\) ms [2, p 118], while we used \(t_{ref} = 2\) ms in 2.

References

1(1,2): Brette R et al. (2007). Simulation of networks of spiking neurons: A review of tools and strategies. Journal of Computational Neuroscience 23:349-98. DOI: https://doi.org/10.1007/s10827-007-0038-6
2(1,2,3,4): Traub RD and Miles R (1991). Neuronal networks of the hippocampus. Cambridge University Press, Cambridge UK.
3: http://modeldb.yale.edu/83319

Parameters

Name	Physical unit	Default value	Description
g_Na	nS	20000nS	Na Conductance
g_K	nS	6000nS	K Conductance
g_L	nS	10nS	Leak Conductance
C_m	pF	200pF	Membrane Capacitance
E_Na	mV	50mV	Reversal potentials
E_K	mV	-90mV	Potassium reversal potential
E_L	mV	-60mV	Leak reversal potential (aka resting potential)
V_T	mV	-63mV	Voltage offset that controls dynamics. For default
tau_syn_exc	ms	5ms	parameters, V_T = -63 mV results in a threshold around -50 mV.Synaptic time constant of excitatory synapse
tau_syn_inh	ms	10ms	Synaptic time constant of inhibitory synapse
t_ref	ms	2ms	Refractory period
E_exc	mV	0mV	Excitatory synaptic reversal potential
E_inh	mV	-80mV	Inhibitory synaptic reversal potential
alpha_n_init	1 / ms	0.032 / (ms * mV) * (15.0mV - E_L) / (exp((15.0mV - E_L) / 5.0mV) - 1.0)
beta_n_init	1 / ms	0.5 / ms * exp((10.0mV - E_L) / 40.0mV)
alpha_m_init	1 / ms	0.32 / (ms * mV) * (13.0mV - E_L) / (exp((13.0mV - E_L) / 4.0mV) - 1.0)
beta_m_init	1 / ms	0.28 / (ms * mV) * (E_L - 40.0mV) / (exp((E_L - 40.0mV) / 5.0mV) - 1.0)
alpha_h_init	1 / ms	0.128 / ms * exp((17.0mV - E_L) / 18.0mV)
beta_h_init	1 / ms	(4.0 / (1.0 + exp((40.0mV - E_L) / 5.0mV))) / ms
I_e	pA	0pA	constant external input current

State variables

Name	Physical unit	Default value	Description
r	integer	0	counts number of tick during the refractory period
V_m	mV	E_L	Membrane potential
Act_m	real	alpha_m_init / (alpha_m_init + beta_m_init)
Act_h	real	alpha_h_init / (alpha_h_init + beta_h_init)
Inact_n	real	alpha_n_init / (alpha_n_init + beta_n_init)

Equations

\[\frac{ dV_{m} } { dt }= \frac 1 { C_{m} } \left( { (-I_{Na} - I_{K} - I_{L} - I_{syn,exc} - I_{syn,inh} + I_{e} + I_{stim}) } \right)\]

\[\frac{ dAct_{m} } { dt }= (\alpha_{m} - (\alpha_{m} + \beta_{m}) \cdot Act_{m})\]

\[\frac{ dAct_{h} } { dt }= (\alpha_{h} - (\alpha_{h} + \beta_{h}) \cdot Act_{h})\]

\[\frac{ dInact_{n} } { dt }= (\alpha_{n} - (\alpha_{n} + \beta_{n}) \cdot Inact_{n})\]

Source codeneuron hh_cond_exp_traub:
  state:
    r integer = 0 # counts number of tick during the refractory period
    V_m mV = E_L # Membrane potential
    Act_m real = alpha_m_init / (alpha_m_init + beta_m_init)
    Act_h real = alpha_h_init / (alpha_h_init + beta_h_init)
    Inact_n real = alpha_n_init / (alpha_n_init + beta_n_init)
  end
  equations:
    # synapses: exponential conductance
    kernel g_inh = exp(-t / tau_syn_inh)
    kernel g_exc = exp(-t / tau_syn_exc)
    # Add aliases to simplify the equation definition of V_m
    inline I_Na pA = g_Na * Act_m * Act_m * Act_m * Act_h * (V_m - E_Na)
    inline I_K pA = g_K * Inact_n * Inact_n * Inact_n * Inact_n * (V_m - E_K)
    inline I_L pA = g_L * (V_m - E_L)
    inline I_syn_exc pA = convolve(g_exc,exc_spikes) * (V_m - E_exc)
    inline I_syn_inh pA = convolve(g_inh,inh_spikes) * (V_m - E_inh)
    V_m'=(-I_Na - I_K - I_L - I_syn_exc - I_syn_inh + I_e + I_stim) / C_m
    # channel dynamics
    inline V_rel mV = V_m - V_T
    inline alpha_n 1/ms = 0.032 / (ms * mV) * (15.0mV - V_rel) / (exp((15.0mV - V_rel) / 5.0mV) - 1.0)
    inline beta_n 1/ms = 0.5 / ms * exp((10.0mV - V_rel) / 40.0mV)
    inline alpha_m 1/ms = 0.32 / (ms * mV) * (13.0mV - V_rel) / (exp((13.0mV - V_rel) / 4.0mV) - 1.0)
    inline beta_m 1/ms = 0.28 / (ms * mV) * (V_rel - 40.0mV) / (exp((V_rel - 40.0mV) / 5.0mV) - 1.0)
    inline alpha_h 1/ms = 0.128 / ms * exp((17.0mV - V_rel) / 18.0mV)
    inline beta_h 1/ms = (4.0 / (1.0 + exp((40.0mV - V_rel) / 5.0mV))) / ms
    Act_m'=(alpha_m - (alpha_m + beta_m) * Act_m)
    Act_h'=(alpha_h - (alpha_h + beta_h) * Act_h)
    Inact_n'=(alpha_n - (alpha_n + beta_n) * Inact_n)
  end

  parameters:
    g_Na nS = 20000nS # Na Conductance
    g_K nS = 6000nS # K Conductance
    g_L nS = 10nS # Leak Conductance
    C_m pF = 200pF # Membrane Capacitance
    E_Na mV = 50mV # Reversal potentials
    E_K mV = -90mV # Potassium reversal potential
    E_L mV = -60mV # Leak reversal potential (aka resting potential)
    V_T mV = -63mV # Voltage offset that controls dynamics. For default
    # parameters, V_T = -63 mV results in a threshold around -50 mV.

    # parameters, V_T = -63 mV results in a threshold around -50 mV.
    tau_syn_exc ms = 5ms # Synaptic time constant of excitatory synapse
    tau_syn_inh ms = 10ms # Synaptic time constant of inhibitory synapse
    t_ref ms = 2ms # Refractory period
    E_exc mV = 0mV # Excitatory synaptic reversal potential
    E_inh mV = -80mV # Inhibitory synaptic reversal potential
    alpha_n_init 1/ms = 0.032 / (ms * mV) * (15.0mV - E_L) / (exp((15.0mV - E_L) / 5.0mV) - 1.0)
    beta_n_init 1/ms = 0.5 / ms * exp((10.0mV - E_L) / 40.0mV)
    alpha_m_init 1/ms = 0.32 / (ms * mV) * (13.0mV - E_L) / (exp((13.0mV - E_L) / 4.0mV) - 1.0)
    beta_m_init 1/ms = 0.28 / (ms * mV) * (E_L - 40.0mV) / (exp((E_L - 40.0mV) / 5.0mV) - 1.0)
    alpha_h_init 1/ms = 0.128 / ms * exp((17.0mV - E_L) / 18.0mV)
    beta_h_init 1/ms = (4.0 / (1.0 + exp((40.0mV - E_L) / 5.0mV))) / ms
    # constant external input current
    I_e pA = 0pA
  end
  internals:
    RefractoryCounts integer = steps(t_ref)
  end
  input:
    inh_spikes nS <-inhibitory spike
    exc_spikes nS <-excitatory spike
    I_stim pA <-current
  end

  output: spike

  update:
    U_old mV = V_m
    integrate_odes()
    # sending spikes: crossing 0 mV, pseudo-refractoriness and local maximum...
    if r > 0:
      r -= 1
    elif V_m > V_T + 30mV and U_old > V_m:
      r = RefractoryCounts
      emit_spike()
    end
  end

end

hh_cond_exp_traub

Description

References

See also

Parameters

State variables

Equations

Source code

Characterisation