The Goldman equation

• The Nernst equation accurately describes the relation

between ion concentrations and the membrane
potential, when only one type of ion can move
through the membrane.
• In reality, many types of ions are present in the intra
and extracellular medium, each of which has its own
permeability.
• For instance, imagine what would happen if the
intracellular K+ and Na+ concentrations are 100 mM
and 10mM, respectively and the extracellular
concentrations of K+ and Na+ are reversed: 10 mM
for K+ and 100 mM for Na+.
• If the membrane would be only permeable to
K+, the membrane potential would be -58 mV.
• If the membrane would be only permeable to
Na+, the membrane potential would be +58 mV.
• What will happen if the membrane is partially
permeable for both ions?
• For the case most relevant to neurons, in which
K+, Na+ and Cl- are the primary permeant ions,
the general solution, called the Goldman
equation, was developed by David Goldman in
1943:
• where V is the equilibrium voltage across the
membrane, and the Pi; i =K+; Na+; Cl- are the
permeabilities of the three types of ions.
• We see that in the case that the membrane is only
permeable to one ion, the Goldman equation
reduces to the Nernst equation.
 The Nernst-Planck equation
• Life is not so simple: no living cell is so
selectively permeable or is in electrochemical
equilibrium.
• The membranes of living cells are invariably
permeable to more than one kind of ion.
• To calculate the membrane potential of such cells,
we have to understand the movement of ions
across cell membranes under non-equilibrium
conditions; and to do that, we have to consider
the forces that act on ions in solution.
• We begin with the same simple system consisting of
two solutions separated by a membrane at constant
pressure and at thermal equilibrium.
• Now we imagine that the solution contains more than
one ion species to which the membrane is permeable.
• What determines the flux of the jth ion (φj) from one
solution to the other?
• There are two forces that act on the ion. One is called
electrophoresis: ions move as charged particles in
response to a difference of potential across the
membrane.
• The other force is diffusion: ions move from a region
of high concentration to one of low concentration.
• If a membrane is placed between two solutions containing
ions at different concentrations, ions will move from one
solution to the other across the membrane, with the total
flux determined by the sum of the electrophoretic and
diffusive fluxes.
• To calculate the net flux, we begin with the component
attributable to electrophoresis (φje ).
• Ions move in an electric field because the field (E)
produces a force, F = qE (where q is the charge).
• The flux of the jth ion is therefore proportional to the
magnitude of the electric field and the amount of charge,
which in turn is determined by the concentration of the
ion in moles per liter (cj) and its valence (zj),
• The flux will, however, also depend on the
mobility of the ion (uj),which quantifies the ease
with which the ion moves across the membrane
in response to an electric field and is determined
ultimately by the size and shape of the ion and
the properties of the membrane.
• The electrical mobility is defined as the ratio of
the velocity of the ion (in cm s−1) to the
amplitude of the electric field.
• We have, then,
• The electric field is equal to the negative of the
gradient of the voltage, or E = −dΨ/dx for a single
dimension across the membrane.
• The flux is negative because ions with positive
valence move from positive to negative, in a direction
opposite to the gradient of potential.
• We can therefore calculate the electrophoretic
component φje of the flux of the jth ion across the
membrane, in units of moles per second per cm2 of
membrane, from the relation
• The second component of flux φjd is produced by
diffusion.
• It is given by Fick’s first law, which states that the
diffusive flux is proportional to the amplitude of
the concentration gradient.
• For a single dimension across the membrane,

• where Dj is the diffusion constant of the jth ion.

The flux is negative because diffusion occurs in a
direction of decreasing concentration from high to
low.
• It is convenient to express the diffusion constant
in terms of the mobility.
• The electrical mobility is related to the diffusion
constant by the Einstein relation, uj = Dje/kT,
where e is the elementary charge and k is
Boltzmann’s constant. Fick’s law is then
• Multiplying both the numerator and the
denominator of by Avogadro’s number, NA, and
remembering that the gas constant R is equal to
kNA and F is equal to eNA, we have

• Summing the diffusional and electrophoretic

components

Nernst Planck eq
• It is helpful to convert this eq. from an eq. of flux
into an equation of current.
• We can do that by multiplying the flux φ j by zjF,
where zj is the valence (or the amount of charge
per ion) and F is the Faraday and gives the
amount of charge per mole.
• We then have the Nernst- Planck equation for the
current density(Ij , expressed in amperes per cm2
of membrane):
• Consider a membrane of thickness a and let x = 0 to x=a
denote the outside and inside of the membrane,
respectively.
• Let the electrical potential be Vout = V (0) = 0 and Vin =
V (a) = V .
• Let us assume that we have a number of different ions,
labeled by i, each with its own concentration Ci(x) and
valence zi.
• The extracellular and intracellular concentrations are
denoted by
• Due to the potential difference and the
concentration differences, ions will flow through
the membrane.
• The electric force per ion of type i is
• The number of ions per liter is N ACi(x), with Ci(x)
in units of mol per liter.
• Therefore, the electric force per unit volume is
• The diffusive force on ion i per unit volume is
proportional to the concentration gradient as well
as the absolute temperature.
• Multiplying by the Gas constant gives the
diffusive force in units of Newton per liter.
• The force on the ions is the sum of the electric and
diffusive force and results in a movement of the
ions proportional to the mobility ui of the ions.
• The mobility will be soon related to the specific
permeability of the membrane to each of the ions.
• The direction of movement is in the direction of the
force, but the direction of charge depends again on
the valence. Therefore, the current for ions i is:

In the equation Ci(x), given the values of Ii and the electric potential
V (x) as a function of x and is known as the Nernst-Planck equation.
The ion concentration and the electric potential depend on location
x, the current is independent of x.
• To solve the Eq., we should also describe how V
(x) depends on the charge distributions C i(x)
using Gauss' law.
• Instead, we will make the simplifying assumption
that the membrane potential changes linearly
from the outside to the inside of the membrane,
dV/dx = V/a for 0 < x < a. Then we can solve
with the boundary conditions

The Eq. predicts the ion current that results from an electrical
potential dierence together with a ionic concentration difference.
• We now consider the case where we have various
ion concentrations inside and outside the cell.
• If i = K+; Na+; Cl- The total current is given by
• Then we can solve for the membrane potential V in
terms of the various concentrations inside and
outside the cell. The solution is given by the
Goldman Eq.

G has units of Ω-1 per liter and is called the conductance

and is the inverse resistance. V0 the equilibrium
membrane potential given by the Goldman equation.
 Sample problem
• Calculate the Nernst potential for a cell at
25°C whose membrane is permeable only to
K+ if the internal K+ concentration [K]i is
100mM and the external K+ concentration [K]o
is 2mM.
• Answer: At 25°C, RT/F is 25.7 mV.