Action potentials are the rapid changes in charge across the membrane that occur when a neuron is firing
- Action potentials occur in three main stages: depolarization, repolarization and a refractory period
Depolarisation refers to a sudden change in membrane potential – usually from a (relatively) negative to positive internal charge
- In response to a signal initiated at a dendrite, sodium channels open within the membrane of the axon
- As Na+ ions are more concentrated outside of the neuron, the opening of sodium channels causes a passive influx of sodium
- The influx of sodium causes the membrane potential to become more positive (depolarisation)
Repolarisation refers to the restoration of a membrane potential following depolarisation (i.e. restoring a negative internal charge)
- Following an influx of sodium, potassium channels open within the membrane of the axon
- As K+ ions are more concentrated inside the neuron, opening potassium channels causes a passive efflux of potassium
- The efflux of potassium causes the membrane potential to return to a more negative internal differential (repolarisation)
The refractory period refers to the period of time following a nerve impulse before the neuron is able to fire again
- In a normal resting state, sodium ions are predominantly outside the neuron and potassium ions mainly inside (resting potential)
- Following depolarisation (sodium influx) and repolarisation (potassium efflux), this ionic distribution is largely reversed
- Before a neuron can fire again, the resting potential must be restored via the antiport action of the sodium-potassium pump
• An action potential consists of depolarization and repolarization of the neuron
Here is the structure of a myelinated motor neurone:
Labelled “insulating sheath”, the myelin sheath is responsible for protecting the electrical impulses that run across the neurone.
But first, what happens in a resting state where no impulses are being sent?
This is the resting potential where the membrane permeability differentiates between sodium (Na+) and potassium (K+) ions so that at any given time there are more Na+ ions outside than inside and more K+ ions inside than outside.
According to these electrochemical gradients, Na+ ions should move back inside to balance out their concentration (equilibrate) while K+ ions should move back outside the membrane until the concentrations are equal inside and out. This clearly isn’t the case, so what gives?
Found on the membrane there are Na+/K+ pumps which carry out active transport against the electrochemical gradient of these ions. The resting potential of the membrane is negative on the inside and positive on the outside – but how? Aren’t both sodium and potassium ions positively charged? This is achieved by the pump transferring 3 Na+ ions out while taking only 2 K+ ions in. This is where the difference comes from.
Now we know that in the absence of an action potential the resting potential of the neurone membrane is negative (about -70mV; millivolts). What precedes an action potential and how does it unfold?
A stimulus may depolarise the membrane by opening up Na+ channels for those ions to rush into the axon. An action potential will occur only if the depolarisation passes a certain threshold. For example, if it reaches -60mV up from -70mV it will not trigger an action potential with a threshold of >-45mV.
Therefore, the power of an action potential is not proportional to that of its stimulus. It either happens or it doesn’t. This is called the all-or-nothing principle.
This is how the voltage of the axon membrane changes during an action potential:
The rush of Na+ ions into the membrane during depolarisation causes the voltage to become positive. Note how only the depolarisation that has passed the threshold initiates an action potential.
Repolarisation occurs when Na+ channels begin to close and K+ channels open, resulting in a rush of K+ ions out of the axon. Before all the K+ channels close, hyperpolarisation occurs which briefly sees the voltage drop below the resting potential level.
This also represents the refractory period where either no stimulus however strong can initiate another action potential (absolute refractory period), or a stimulus slightly greater than usual would be required for an action potential to occur (relative refractory period).
Finally the resting potential is achieved.
The Myelin Sheath
This insulating sheath made up of Schwann cells conducts electricity and therefore is key in ensuring fast signal transmission. The signal is able to “jump” along the axon without losing its strength:
Each pink cell is a Schwann cell. Due to the jump-like action, this conduction is termed saltatory conduction. Factors that affect conduction other than myelination and saltatory conduction (which allow speeds many times faster compared with no myelination) include temperature and axon diameter.
Since chemical movement (kinetic energy) relies on temperature, an optimal temperature maximises conduction. A temperature lower than this would slow it down. This is due to a slower opening of sodium channels for example, and also a slower inactivation resulting in a longer delay.
Axon diameter affects conduction in terms of resistance. The signal travelling along a thin axon encounters the resistance of the axon membrane, while for an axon with larger diameter, a smaller proportion of the signal is met with resistance in this way. The signal carried on the inner section of the axon has no resistance and can travel faster.