The membrane is a capacitor () pierced by ion channels in parallel. Conservation of charge (Kirchhoff) balances the capacitive current against the total ionic current:
Each channel passes an ohmic current — its conductance times the driving force , the distance of from the reversal potential (where the channel’s net current vanishes, set by the Nernst equilibrium):
Because , the current’s sign lives entirely in the driving force. Summing a leak (, ) and synaptic conductances — excitatory (, ), inhibitory (, ) — gives the general conductance-based (COBA) neuron:
COBANet specialises (3) to two populations — E driven by excitation and inhibition, I by excitation only:
A neuron spikes at threshold and resets to for a refractory period:
Each synaptic conductance is an exponential trace driven by presynaptic spikes — each spike adds its full weight as an instantaneous jump, then the conductance decays with the channel time constant; there is no E→E connection:
(7) is E’s excitation from the input ; (8) its inhibition from I via ; (9) the I population’s excitation from E via .
The conductances (7)–(9) and membrane equations (4)–(5) are continuous ODEs. The delta-driven conductances integrate exactly over one step: between spikes they decay by , and any spike landing in the step adds its full weight — the decay-then-add recurrence (with the , and spike train of each of (7)–(9)). The membrane we integrate by exponential Euler — the same algebra for both populations (the I neuron drops ).
Collecting on makes it linear, with total conductance :
Dividing by gives decay-to-steady-state form, naming (shorter than when synapses are open) and the steady-state voltage (the conductance-weighted mean of the reversals):
A zero-order hold freezes the conductances over one step , leaving (11) constant-coefficient with exact solution
Per population — I has no , so its and drop those terms:
with step (12) for each population : .
Being the exact frozen-conductance integral, (12) is dt-invariant — small steps equal one big step, so firing rates and the gamma frequency are physical (Hz) properties, not timestep artifacts (exp044). A forward-Euler step — not dt-invariant — is kept only as a parity toggle (COBA_INTEGRATOR).
Each step runs in fixed order: conductances (7)–(9), then , then the membrane step (12), then spike + reset (6). The zero-order hold is this ordering — the conductances advance once, then stay fixed while the membrane integrates across . E and I advance synchronously, phase-locking the E→I→E gamma cycle to the grid.