Supplement: Electrical Characteristics of Neurons

Goldman Equation vs. Parallel Conductance Model

The Goldman-Hodgkin-Katz equation (GHK) shows the dependence of Vm on
concentrations and permeabilities:

PK [K + ]out + PNa [Na + ]out + PCl [Cl" ]in

PK [K + ]in + PNa [Na + ]in + PCl [Cl" ]out
GHK does not directly predict the flux of ions, which depends not only on permeability
but also on absolute concentration.
The Parallel Conductance Model (PCM) of the membrane gives a different way of
!
calculating
Vm, based on conductivity and ion potentials:
g E + g NaE Na + g ClE Cl
Vm = K K
g K + g Na + g Cl
This comes from an electrical model of the membrane and is easily derived from basic
physics and algebra (Ohms law and Kirchoffs law). See next page if interested.
Vm = 58log

! Conductance
Permeability and
Permeability and conductance are related but not the same. Permeability is the ease with
which an ion can pass through the membrane. Conductance is the ability of a given ion
to carry electrical current across the membrane.
Conductance depends on concentration as well as permeability. For example,
permeability could be high (many open channels) but if only a few ions are present,
then that ion species cant carry much current.
Conductance and current are easier to measure than permeability and ion movement.
Currents are measured and conductances calculated in voltage or patch clamp.
Measuring permeability requires some way of observing ion movement itself.
In practice, neurophysiologists generally use the PCM method because it involves
parameters that are more readily available. Note, too, that we can measure Eion directly
or calculate it from other electrical measures, but we rarely have direct access to
[ion]in. That is another reason to use PCM instead of GHK.
The exact values of the permeabilities (or conductances) do not matter. All that matters is
their ratios. See this by simple algebraic rearrangement:

Vm = 58log

[K + ]out +
+

[K ]in +

PNa
PK
PNa
PK

[Na + ]out +
+

[Na ]in +

PCl
PK

PCl
PK

[Cl" ]in
"

[Cl ]out

Vm =

EK +

g Na
gK

1+

E Na +
g Na
gK

g Cl
gK

E Cl

g Cl
gK

Dividing by PK or gK is arbitrary (mathematically, it works out the same if we choose

another ion instead), but it is usual to divide by the permeability or conductance that is
greatest at rest. If the permeability (or conductance) to one ion is much smaller than
!
for the others, the term for that ion may be omitted.
Bottom line: use conductance and the PCM equation unless permeability is specified.
Reversal Potential
Electromotive force (driving force) for an ion = Vm Eion. This is the force that drives
the ion through open channels. The amount of ion current is gion(Vm Eion). If driving
force is zero, the ion does not move. If conductance is zero, the ion does not move.
Reversal potential (Erev) is the membrane potential at which the sign of a current reverses
(e.g. a current that is inward at RP becomes outward above Erev).
Erev is simply the point where the driving force is zero. Thus for a single ion, it is Eion
(when Vm = Eion, there is no driving force). When Vm < Eion, Iion is negative, indicating
an inward current. When Vm > Eion, Iion is positive, indicating an outward current.

We often speak of the reversal potential of a channel. For a simple ion channel, Erev is the
same as the equilibrium potential of the ion that it passes. When the channel opens, it
brings Vm toward the equilibrium potential for that ion.
For a channel that allows more than one ion to pass, Erev is more complex, depending on
the conductivity to and equilibrium potential of each ion.
Example: the acetylcholine receptor is a ligand-gated ion channel equally permeable to
Na+ and K+. Its reversal potential should therefore be halfway between EK and ENa
(using the PCM equation). When the acetylcholine receptor channel opens, it brings
Vm toward the reversal potential (often around 0mV).
Why do we care about reversal potential? Because it is relatively easy to measure
experimentally. It is often the first thing we know about a channel, before we know
anything about what ion(s) it passes. In fact, it is through knowing Erev and Eion for all
the relevant ions that we get a clue about what ions the channel can pass.
Reminder About Ion Movement
At electrochemical equilibrium, very few ions have crossed the membrane. (Its very few
ions by standards of mM concentrations; its actually something like 1012 ions.)
Those ions that do move stay close to the membrane.
Thus the ion concentrations, for all practical purposes, do not change.
Thus each compartment (inside the cell, outside the cell) remains electrically neutral.
That is, there is no net charge inside the cell; its cations and anions are balanced.
Of course, some ions do move, and the cell has active ion transporters to maintain its ion
gradients. Without these, the gradients would eventually disappear and Vm = 0 (dead).
Parallel Conductance Model Derivation
(You dont need to worry about this, but heres the simple derivation I promised.)
Kirchoffs law states that the sum of all currents entering and leaving a node is zero.
Ohms law states that V = IR (or I = gV, since g = 1/R).
From Kirchoffs law, we know that IK + INa + ICl Im = 0.
At rest, Im is zero. Thats what it means for the membrane to be at rest.
From Ohms law, Iion = gion(Vm Eion). For V, we use the driving force for each ion.
Substitute gion(Vm Eion) for each of IK, INa, and ICl above:
gK(Vm EK) + gNa(Vm ENa) + gCl(Vm ECl) = 0
multiply the conductances through and rearrange:
Vm(gK + gNa + gCl) (gKEK + gNaENa + gClECl) = 0
now solve for Vm:
g E + g NaE Na + g ClE Cl
Vm = K K
.
g K + g Na + g Cl