Squid giant axon
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The squid giant axon is the very large (up to 1 mm in diameter; typically around 0.5 mm) axon that controls part of the water jet propulsion system in squid. It was first described by L. W. Williams in 1909, but this discovery was forgotten until English zoologist and neurophysiologist J. Z. Young demonstrated the axon’s function in the 1930s while working in Naples (Stazione Zoologica), Plymouth (the Marine Biological Association) and Woods Hole (Marine Biological Laboratory). Squids use this system primarily for making brief but very fast movements through the water.
Between the tentacles of a squid is a siphon through which water can be rapidly expelled by the fast contractions of the body wall muscles of the animal. This contraction is initiated by action potentials in the giant axon. Action potentials travel faster in a larger axon than a smaller one, and squid have evolved the giant axon to improve the speed of their escape response. The increased diameter of the squid axon decreases the internal resistance of the axon, as resistivity is inversely proportional to the cross sectional area of the object. This increases the space constant, λ=sqrt(rm/ri).
The increased space constant propagates a given local depolarization further, which speeds up the action potential, according to the equation
In their Nobel Prize-winning work uncovering ionic mechanism of action potentials, Alan Hodgkin and Andrew Huxley performed experiments on the squid giant axon. The prize was shared with John Eccles. The large diameter of the axon provided a great experimental advantage for Hodgkin and Huxley as it allowed them to insert voltage clamp electrodes inside the lumen of the axon.
While the squid axon is very large in diameter it is unmyelinated which decreases the conduction velocity substantially. The conduction velocity of a typical 0.5 mm squid axon is about 25 m/s. During a typical action potential in the cuttlefish Sepia giant axon, an influx of 3.7 pmol/cm2 (picomoles per centimeter2) of sodium is offset by a subsequent efflux of 4.3 pmol/cm2 of potassium.
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