The Hodgkin–Huxley model, or conductance-based model, is a mathematical model that describes how action potentials in neurons are initiated and propagated. It is a set of nonlinear differential equations that approximates the electrical engineering characteristics of excitable cells such as neurons and muscle cells. It is a continuous-time dynamical system. Alan Hodgkin and Andrew Huxley described the model in 1952 to explain the ionic mechanisms underlying the initiation and propagation of action potentials in the squid giant axon. They received the 1963 Nobel Prize in Physiology or Medicine for this work.
Basic components The typical Hodgkin–Huxley model treats each component of an excitable cell as an electrical element (as shown in the figure). The lipid bilayer is represented as a capacitance (Cm). Voltage-gated ion channels are represented by electrical conductances (gn, where n is the specific ion channel) that depend on both voltage and time. Leak channels are represented by linear conductances (gL). The electrochemical gradients driving the flow of ions are represented by voltage sources (En) whose voltages are determined by the ratio of the intra- and extracellular concentrations of the ionic species of interest. Finally, ion pumps are represented by current sources (Ip). The membrane potential is denoted by Vm. Mathematically, the current flowing into the capacitance of the lipid bilayer is written as
I c = C m d V m d t {\displaystyle I_{c}=C_{m}{\frac {{\mathrm {d} }V_{m}}{{\mathrm {d} }t}}}
and the current through a given ion channel is the product of that channel's conductance and the driving potential for the specific ion
I i = g i ( V m − V i ) {\displaystyle I_{i}={g_{i}}(V_{m}-V_{i})\;}
where V i {\displaystyle V_{i}} is the reversal potential of the specific ion channel. Thus, for a cell with sodium and potassium channels, the total current through the membrane is given by:
I = C m d V m d t + g K ( V m − V K ) + g N a ( V m − V N a ) + g l ( V m − V l ) {\displaystyle I=C_{m}{\frac {{\mathrm {d} }V_{m}}{{\mathrm {d} }t}}+g_{K}(V_{m}-V_{K})+g_{Na}(V_{m}-V_{Na})+g_{l}(V_{m}-V_{l})}
where I is the total membrane current per unit area, Cm is the membrane capacitance per unit area, gK and gNa are the potassium and sodium conductances per unit area, respectively, VK and VNa are the potassium and sodium reversal potentials, respectively, and gl and Vl are the leak conductance per unit area and leak reversal potential, respectively. The time dependent elements of this equation are Vm, gNa, and gK, where the last two conductances depend explicitly on the membrane voltage (Vm) as well.
Ionic current characterization In voltage-gated ion channels, the channel conductance is a function of both time and voltage ( g n ( t , V ) {\displaystyle g_{n}(t,V)} in the figure), while in leak channels, g l {\displaystyle g_{l}} , it is a constant ( g L {\displaystyle g_{L}} in the figure). The current generated by ion pumps is dependent on the ionic species specific to that pump. The following sections will describe these formulations in more detail.
Voltage-gated ion channels Using a series of voltage clamp experiments and by varying extracellular sodium and potassium concentrations, Hodgkin and Huxley developed a model in which the properties of an excitable cell are described by a set of four ordinary differential equations. Together with the equation for the total current mentioned above, these are:
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