COBANet

COBANet is a conductance-based leaky integrate-and-fire (LIF) spiking neural network with separate excitatory (E) and inhibitory (I) populations, fixed recurrent E↔I connectivity, and exponential synapses. This article derives the model equations and states the architectural constants. All parameter values follow Börgers (2017).

Neuron model

Membrane dynamics

Each neuron’s membrane potential VV evolves according to the conductance-based LIF equation:

CmdVdt=gL(VEL)ge(VEe)gi(VEi)(1)C_m\, \frac{dV}{dt} = -g_L(V - E_L) - g_e(V - E_e) - g_i(V - E_i) \tag{1}

where CmC_m is capacitance, gLg_L is leak conductance, geg_e and gig_i are the total excitatory and inhibitory synaptic conductances, and ELE_L, EeE_e, EiE_i are the corresponding reversal potentials. Each term pulls VV toward its reversal at a rate proportional to the conductance.

Collecting on VV:

CmdVdt=(gL+ge+gi)V+(gLEL+geEe+giEi)(2)C_m\, \frac{dV}{dt} = -(g_L + g_e + g_i)\, V + (g_L E_L + g_e E_e + g_i E_i) \tag{2}

Define the total conductance, effective time constant, and steady-state voltage:

gtot=gL+ge+gi,τeff=Cmgtot,V=gLEL+geEe+giEigtot(3)g_{\text{tot}} = g_L + g_e + g_i, \qquad \tau_{\text{eff}} = \frac{C_m}{g_{\text{tot}}}, \qquad V_\infty = \frac{g_L E_L + g_e E_e + g_i E_i}{g_{\text{tot}}} \tag{3}

Then (2) becomes:

τeffdVdt=(VV)(4)\tau_{\text{eff}}\, \frac{dV}{dt} = -(V - V_\infty) \tag{4}

Note that τeff\tau_{\text{eff}} is not the leak-only time constant τm=Cm/gL\tau_m = C_m / g_L — when synapses are open, gtot>gLg_{\text{tot}} > g_L and the membrane responds faster. This is shunting inhibition: open gig_i increases gtotg_{\text{tot}}, shortening τeff\tau_{\text{eff}} and dampening the membrane’s response to all inputs.

Discretisation

Hold geg_e, gig_i constant over [t,t+Δt][t, t + \Delta t] (zero-order hold). Then τeff\tau_{\text{eff}} and VV_\infty are constant and (4) integrates exactly:

Vt+1=V+(VtV)exp ⁣(Δt/τeff)(5)V_{t+1} = V_\infty + (V_t - V_\infty)\,\exp\!\left(-\Delta t / \tau_{\text{eff}}\right) \tag{5}

Spike and reset

St+1=1[Vt+1Vth],Vt+1Vreset  if  St+1=1  or refractory(6)S_{t+1} = \mathbf{1}[V_{t+1} \geq V_{\text{th}}], \qquad V_{t+1} \leftarrow V_{\text{reset}} \;\text{if}\; S_{t+1} = 1 \;\text{or refractory} \tag{6}

with Vth=50V_{\text{th}} = -50 mV and Vreset=65V_{\text{reset}} = -65 mV. Reset is hard (overshoot discarded). After spiking the neuron enters a refractory period during which VV is clamped at VresetV_{\text{reset}} and no spike can be emitted.

Synaptic conductances

Each conductance is the convolution of its presynaptic spike train with a decaying exponential:

ge,t+1=αge,t+Wst,α=eΔt/τAMPA(7a)g_{e,\,t+1} = \alpha\, g_{e,\,t} + W\, s_t, \qquad \alpha = e^{-\Delta t / \tau_{\text{AMPA}}} \tag{7a} gi,t+1=γgi,t+Wst,γ=eΔt/τGABA(7b)g_{i,\,t+1} = \gamma\, g_{i,\,t} + W\, s_t, \qquad \gamma = e^{-\Delta t / \tau_{\text{GABA}}} \tag{7b}

Between spikes the conductance decays exponentially toward zero; each presynaptic spike adds a kick proportional to the synaptic weight WW. Excitatory synapses use the AMPA time constant (τAMPA=2\tau_{\text{AMPA}} = 2 ms); inhibitory synapses use the GABA time constant (τGABA=9\tau_{\text{GABA}} = 9 ms).

Population structure

The network has a single hidden layer partitioned into E and I sub-populations in a 4:1 ratio. E and I neurons obey the same LIF equations (1)–(6) but with different biophysical constants:

propertysymbolEI
membrane time constantτm\tau_m20 ms5 ms
capacitanceCmC_m1.0 nF0.5 nF
leak conductancegL=Cm/τmg_L = C_m / \tau_m0.05 µS0.1 µS
refractory periodτref\tau_{\text{ref}}3 ms1.5 ms
synaptic decay onto targetτsyn\tau_{\text{syn}}2 ms (AMPA)9 ms (GABA)
reversal at targetEe=0E_e = 0 mVEi=80E_i = -80 mV
population fraction4/51/5

Three asymmetries matter for the PING cycle:

  1. I neurons integrate four times faster (τmI=5\tau_m^I = 5 ms vs τmE=20\tau_m^E = 20 ms). A synchronous E-burst raises I above threshold within a single AMPA window (≈2 ms).
  2. GABA decay is 4.5× slower than AMPA (τGABA=9\tau_{\text{GABA}} = 9 ms vs τAMPA=2\tau_{\text{AMPA}} = 2 ms). The resulting I→E inhibition outlasts the triggering excitation, suppressing E long enough for excitatory drive to re-accumulate. This sets the gamma period.
  3. I neurons receive no inhibition. Only geg_e is wired into the I update. The I population follows E’s drive without self-suppression.

Recurrent connectivity

There are no E→E connections (WEE=0W^{EE} = 0, Börgers-style PING). Two recurrent pathways close the loop alongside feedforward input:

ge,t+1E=αge,tE+Winstinp(8)g^{E}_{e,\,t+1} = \alpha\, g^{E}_{e,\,t} + W_{\text{in}}\, s^{\text{inp}}_t \tag{8} gi,t+1E=γ(gi,tE+WIEstI)(9)g^{E}_{i,\,t+1} = \gamma\bigl(g^{E}_{i,\,t} + W^{IE}\, s^{I}_t\bigr) \tag{9} ge,t+1I=α(ge,tI+WEIstE)(10)g^{I}_{e,\,t+1} = \alpha\bigl(g^{I}_{e,\,t} + W^{EI}\, s^{E}_t\bigr) \tag{10}

Eq (8): excitatory conductance on E neurons, purely feedforward. Eq (9): inhibitory conductance on E neurons, driven by I spikes through WIEW^{IE}. Eq (10): excitatory conductance on I neurons, driven by E spikes through WEIW^{EI}.

WEIW^{EI} and WIEW^{IE} are controlled by the scalar —ei-strength ss:

WEIN(s,  0.1s),WIEN(rs,  0.1rs)W^{EI} \sim \mathcal{N}(s,\; 0.1\,s), \qquad W^{IE} \sim \mathcal{N}(r\,s,\; 0.1\,r\,s)

where rr is the —ei-ratio (default 2.0). At s=0s = 0 both matrices are zero, the loop is open, and the network is feedforward COBA. At s=1s = 1 the initialisation gives WEI1W^{EI} \approx 1 µS and WIE2W^{IE} \approx 2 µS — sufficient to sustain PING from the first forward pass. Both matrices are held requires_grad = False; the optimiser cannot modify them.

Trainable parameters

Only two weight matrices train:

  • WinW_{\text{in}} (Nin×NEN_{\text{in}} \times N_E): feedforward input to E conductance. Initialised from N(μ,0.1μ)\mathcal{N}(\mu, 0.1\mu) with 95% sparsity. Default μ\mu depends on configuration: 0.3 for COBA, 1.2 for PING.
  • WoutW_{\text{out}} (NE×NclassesN_E \times N_{\text{classes}}): linear readout from the time-averaged E spike vector (mem-mean mode).

All recurrent weights (WEIW^{EI}, WIEW^{IE}), all biophysical constants (τm\tau_m, CmC_m, gLg_L, τAMPA\tau_{\text{AMPA}}, τGABA\tau_{\text{GABA}}, reversal potentials), and the E→E matrix (WEE=0W^{EE} = 0) are frozen.

Constants

constantvalue
ELE_L (leak reversal)65-65 mV
EeE_e (excitatory reversal)00 mV
EiE_i (inhibitory reversal)80-80 mV
VthV_{\text{th}} (spike threshold)50-50 mV
VresetV_{\text{reset}}65-65 mV
τAMPA\tau_{\text{AMPA}}22 ms
τGABA\tau_{\text{GABA}}99 ms
E:I ratio4:1
—ei-ratio rr2.0
Δt\Delta t (recommended)1\leq 1 ms