Propagation of electrical signals along giant nerve fibres

Abstract

In this part of the discussion we shall attempt to describe the way in which electrical signals are propagated along the giant nerve fibres of squids and cuttlefish. These fibres consist of cylinders of protoplasm, 0.2 to 0.6 mm in diameter, and ire bounded by a thin membrane which acts as a barrier to ionic movement. The protoplasm, or axoplasm as it is commonly called, is an aqueous gel which is a reasonably good conductor of electricity. It contains a high concentration of K + and a low concentration of Na + and Cl - , this situation being the reverse of that in the animal’s blood or sea water. Axons which are left in sea water slowly lose potassium and gain sodium. This process takes about 24 hours and is roughly 80 000 times slower than the diffusion of ions out of a cylinder of gelatin of the same size. The interchange of sodium and potassium is very greatly accelerated by stimulating the fibres. Experiments with tracers, such as those made by Keynes & Lewis (1951) or Rothenberg (1950), allow the interchange to be measured quantitatively, and there is general agreement that the impulse is associated with an entry of 3 to 4 µ µ mol of Na + through 1 cm 2 of membrane and an exit of a corresponding quantity of K + . These quantities are very small compared with the total number of ions inside the fibre. In the giant axon of the squid the quantity of potassium lost in each impulse corresponds to only about 1 millionth if the total internal potassium. One would therefore expect that a giant fibre should be able to carry a great many impulses without recharging its batteries by metabolism. On the other hand, a very small fibre such as a dendrite in the central nervous system should be much more dependent on metabolism since the ratio of surface to volume may be nearly 1000 times greater.