Hodgkin, Huxley, and Katz (3) and Hodgkin and Huxley (14), (15), (16),
in 1952, published a series of papers describing detailed measurements
of voltage, current, and time relationships in the giant axon of the
squid (_Loligo_). Hodgkin and Huxley (17) consolidated and formalized
these data into a set of simultaneous differential equations describing
the hypothetical time course of events during spike generation and
propagation. The hypothetical system which these equations describe is
the basis of the Modern Ionic Hypothesis.
The system proposed by Hodgkin and Huxley is basically one of dynamic
opposition of ionic fluxes across the axon membrane. The membrane
itself forms the boundary between two liquid phases—the intracellular
fluid and the extracellular fluid. The intracellular fluid is rich in
potassium ions and immobile organic anions, while the extracellular
fluid contains an abundance of sodium ions and chloride ions. The
membrane is slightly permeable to the potassium, sodium, and chloride
ions; so these ions tend to diffuse across the membrane. When the
axon is inactive (not propagating a spike), the membrane is much more
permeable to chloride and potassium ions than it is to sodium ions.
In this state, in fact, sodium ions are actively transported from the
inside of the membrane to the outside at a rate just sufficient to
balance the inward leakage. The relative sodium ion concentrations on
both sides of the membrane are thus fixed by the active transport rate,
and the net sodium flux across the membrane is effectively zero. The
potassium ions, on the other hand, tend to move out of the cell; while
chloride ions tend to move into it. The inside of the cell thus becomes
negative with respect to the outside. When the potential across the
membrane is sufficient to balance the inward diffusion of chloride with
an equal outward drift, and the outward diffusion of potassium with an
inward drift (and possibly an inward active exchange), equilibrium is
established. The equilibrium potential is normally in the range of 60
to 65 millivolts.
Public-domain text, read in full here on John Shaqi.
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