Conduction Of Action Potentials
The action potential described in
Chapter 5 is a local event that can
occur in all excitable cells. This local event is an all-or-nothing response,
leading to abolishion and then reversal of the polarity from negative (−70 mV)
to positive (+40 mV) on the inside of the cell with respect to the outside for
a short time during the course of the action potential.
Local currents are set up around
the action potential because the positive charges from the membrane ahead of
the action potential are drawn towards the area of negativity surrounding the
action potential (current sink). This decreases the polarity of the membrane
ahead of the action potential.
This electronic depolarization
initiates a local response that causes the opening of the voltage-gated ion
channels (Na+ followed by K+); when the threshold for firing of the action
potential is reached, it propagates the action potential and this, in
turn, leads to the local depolarization of the next area, and so on. Once
initiated, an action potential does not depolarize the area behind it
sufficiently to initiate another action potential because the area is refractory
(Chapter 5).
This successive depolarization
moves along each segment of an unmyelinated nerve until it reaches the
end. It is all-or-nothing and does not decrease in size (Fig. 6a).
Saltatory conduction
Conduction in myelinated axons
depends on a similar pattern of current flows. However, because myelin is
an insulator and because the membrane below it cannot be depolarized, the only
areas of the myelinated axon that can be depolarized are those that are devoid
of any myelin, i.e. at the nodes of Ranvier. The depolarization jumps
from one node to another and is called saltatory, from the Latin saltare
(to jump) (Fig. 6b). Saltatory conduction is rapid and can be up to 50
times faster than in the fastest unmyelinated fibres.
Saltatory conduction not only
increases the velocity of impulse transmission by causing the depolarization
process to jump from one node to the next, but also conserves energy for
the axon because depolarization only occurs at the nodes and not along the
whole length of the nerve fibre, as in unmyelinated fibres. This leads to up to
100 times less movement of ions than would otherwise be necessary, therefore
conserving the energy required to re-establish the Na+ and K+ concentration
differences across the membranes following a series of action potentials being
propagated along the fibre.
All nerve fibres are capable of
conducting impulses in either direction if stimulated in the middle of their
axon; however, normally they conduct impulses in one direction only (orthodromically),
from either the receptor to the axon terminal or from the synaptic junction to
the axon terminal. Antidromic conduction does not normally occur.
Fibre diameters
and conduction velocities Some information needs to be transmitted to and from the central nervous
system very rapidly, whereas other information does not. Nerve fibres are able
to cover both of these extremes and any in between by virtue of their size, and therefore conduction velocity, and
whether or not they are myelinated. Nerve
fibres come in all sizes, from 0.5 to 20 μm in diameter, with the smallest
diameter unmyelinated fibres being the slowest conducting and the largest
myelinated fibres the fastest conducting.
Classification of nerve fibres
Unfortunately, there are two
classifications of nerve fibres. One, originally described by Erlanger and
Gasser, and often referred to as the general classification, uses the
letters A, B and C, with A further subdivided into α, β, γ and
δ. The second, originally described by Lloyd and Hunt, and often referred
to as the sensory or afferent clas- sification, uses the Roman
numerals I, II, III and IV, with I further subdivided into A and B. The
groups are subdivided differently in the two classifications and so,
unfortunately, it is not possible to rely on only one of the classifications
for the description of nerve fibres. The fibres of groups A and B and also
of groups I, II and III are all myelinated, and those of group C and IV are
unmyelinated. These classifications, conduction velocities, fibre diameters
and examples of their functions are shown in Figure 6c. A word of caution is
necessary concerning the average conduction velocities of the larger myelinated
fibres: in reality, although there may be a few larger diameter fibres in the
human body that do indeed conduct impulses as fast as 120/ms, a more common
observation is that the fastest proprioceptive (sensory) fibres conduct at
below 100 m/s, with the average being closer to 80 m/s. The same applies to the
α-motor neurones in that the conduction velocities rarely exceed 90 m/s, with
the average being closer to 60 m/s.
Compound action potentials
Peripheral nerves in most animals
comprise a number of axons bound together by a fibrous tissue called the epineurium.
When extracellular recording electrodes are placed close to a peripheral nerve,
the recorded voltage signal, when an action potential is initiated in the
bundle, is much smaller (microvolts) than that recorded by an electrode
inserted directly into the axon (millivolts). The extracellular recorded signal
is made up of the electrical events occurring in all of the active fibres
within the nerve bundle. If all the nerve fibres in a nerve bundle are
synchronously stimulated at one end of a nerve, and recording electrodes are
placed at a number of locations along the length of the bundle, a compound
action potential is recorded at each electrode. The waveform recorded from
each of the electrodes will differ due to the different conduction velocities
of each group of fibres that makes up the bundle. Theoretically, if the nerve
bundle were to contain examples of all classifications of nerve fibres (i.e.
Aα, Aβ, Aγ, Aδ, B and C), the recorded compound action potential would be seen
as a multi-peaked display, as the action potentials in the fastest conducting
fibres (Aα) would reach the electrode before those in the slowest conducting
fibres (C). Action potentials in the fibres with conduction velocities between
these two extremes would arrive between
these two times (Fig. 6d).
