The Goldman-Hodgkin-Katz equation calculates the electric potential across a membrane due to ion flow through pores.In 1943, David Goldman, a young American physiologist, published an equation that became the cornerstone of modern electrophysiology. Later, Alan Hodgkin and Bernard Katz, working with the giant squid axon, experimentally confirmed its applicability and used it to build a model of the nerve impulse. The Goldman equation looks like this: V_m = (RT/F) * ln((P_K[K⁺]_out + P_Na[Na⁺]_out + P_Cl[Cl⁻]_in) / (P_K[K⁺]_in + P_Na[Na⁺]_in + P_Cl[Cl⁻]_out)). Here, P_K, P_Na, P_Cl are the permeabilities for the respective ions. Note that for chloride, the concentrations in the numerator and denominator are swapped because chloride is a negative ion, and its movement creates a current in the opposite direction. The equation is derived from the assumption of a constant electric field within the membrane (Goldman's condition) and describes the steady-state potential when the net current across the membrane is zero. It is an elegant generalization of the Nernst equation for multiple ions, explaining why the cell doesn't 'choose' a single ion but finds a compromise.
David Goldman derived the equation in 1943 while working at the U.S. Naval Medical Research Institute. He sought a way to describe the electrical properties of biological membranes without resorting to complex models with fixed charges. Alan Hodgkin and Bernard Katz in 1949 applied the Goldman equation to analyze the resting potential of the squid axon and showed that it accurately predicted the potential's dependence on external potassium concentration. Later, Hodgkin and Andrew Huxley used the GHK equation as the basis for their model of action potential generation, for which they received the Nobel Prize in 1963. In the 1950s, the equation was extended to account for divalent ions and active transport.
The Goldman-Hodgkin-Katz equation is used to calculate the resting potential and reversal potential of ionic currents in neurons, muscle cells, and sensory cells. In experiments, ion concentrations and relative permeabilities are measured, then V_m is computed and compared with microelectrode measurements. Limitations: the equation assumes a steady state (no concentration changes over time), a constant field within the membrane, and independent ion movement. It fails at high concentrations when ions interact, and in the presence of electrogenic pumps that generate additional current. For non-steady-state conditions, the full Hodgkin-Huxley system of equations is used.
Calculation of membrane potential in neurons and cardiomyocytes under normal and pathological conditions. Used in drug development targeting ion channels and in neural interfaces.
The giant squid axon, on which the equation was tested, has a diameter of up to 1 mm—hundreds of times thicker than a human neuron. This allowed researchers in the 1940s to insert electrodes and measure the potential directly, which was impossible with thin cells.
| V | variable | potential energy of electron — potential energy joule |
| z | variable | distance from the metal surface — position metre |
| φ | variable | work function of the metal — work function joule |
| F | variable | applied electric field strength — electric field volt per metre |
| e | constant | charge of a single proton — elementary charge = 1.602176634e-19 coulomb |
| ε0 | constant | ability of vacuum to permit electric field lines — vacuum permittivity = 8.8541878128e-12 farad per metre |



