Calculates the equilibrium potential for a single ion. R is the gas constant, T the absolute temperature (in Kelvin, K), z the ion's valence, and F Faraday's constant.
The Membrane Potential (Vm) considers multiple ions simultaneously. Note that Cl− concentrations are inverted due to its negative valence.
Driving Force
DF=Vm−Eion
Determines the net direction and magnitude of an ion's flow across the membrane. For cations, a negative sign indicates the force pushes it into the cell (influx), while a positive sign indicates efflux.
GHK Model Limitations
Active Transport Ignored: The GHK equation describes a passive steady state. It does not explicitly incorporate the small electrogenic flux of the Na⁺/K⁺-ATPase pump (which typically contributes an additional -2 to -5 mV to the Vm).
Ideal Solutions: It assumes that ionic activities are equal to their molar concentrations, ignoring activity coefficients in highly concentrated solutions.
Constant Field: The mathematical derivation assumes that the electric field across the membrane is uniform and constant, which is an approximation (Goldman's constant-field theory).
1. Resting Potential
Living cells maintain a voltage difference across their cell membrane. This is due to the unequal distribution of ions (mainly Na+, K+, and Cl−) and selective permeability. At rest, the membrane is much more permeable to Potassium, so the Vm approaches EK.
2. Nernst vs. GHK
While Nernst assumes permeability to a single type of ion, the GHK equation calculates the potential taking into account the relative permeabilities (P) of all ions simultaneously. If one ion's permeability overwhelmingly dominates, GHK mathematically collapses into Nernst for that ion.
3. Role of Permeability
Ion concentrations change slowly. However, permeability (P) changes in milliseconds by opening channels. In an action potential, PNa increases drastically, causing the Vm to shoot up towards the Sodium equilibrium potential.
4. The Calcium Case (Ca²⁺)
The GHK equation assumes monovalent ions (z=±1). Calcium is divalent (z=+2) and cannot be directly summed. For different valences, the complex Extended GHK Equation is required. Therefore, Calcium's contribution is analyzed by calculating its Nernst independently to understand its Driving Force.
Goldman, D. E. (1943). Potential, Impedance, and Rectification in Membranes. Journal of General Physiology, 27(1), 37-60.
Hodgkin, A. L., & Katz, B. (1949). The effect of sodium ions on the electrical activity of the giant axon of the squid. The Journal of Physiology, 108(1), 37-77.