Action Potential

Ion Channels
- Passive (non-gated) channels are open at all times, permitting ions to move across the
membrane.
- Voltage-gated channels contain a voltage sensitive string of amino acids that cause the
channel pore to open or close in response to changes in membrane voltage
- Channel pumps are energy driven ion exporters and/or importers designed to maintain
steady-state ion concentrations. The Na-K exchange pump (usually referred to as sodium
pump) is vital to maintenance of the resting membrane potential
- Transmitter-gated channels abound in postsynaptic membranes. Some are activated
directly by transmitter molecules, others indirectly.
- Transduction channels are activated by peripheral sensory stimulation. Sensory nerve
endings exhibit different stimulus specificities in different location, for example mechanical
in muscle; tactile, thermal, or chemical in skin; acoustic in cochlea; vestibular in labyrinth;
electromagnetic in the retina; gustatory in the tongue; olfactory in the upper part of the
nasal mucous membrane.
Resting Membrane Potential
- When a neuron is not sending a signal, it is at rest. When a neuron is at rest, the inside
of the neuron is negative relative to the outside.
- The membrane potential of the resting neuron is generated primarily by differences in
concentration of Na and K ions dissolved in the aqueous environments of ECF and cytosol
- The membrane potential ranges from -60mV to -80mV.
Resting Membrane Permeability
K+ Ions
- The K concentration on either side of the cell membrane would be the same if there were
no attraction exerted by the protein anions on the inside, and the repulsion exerted by the
Na cations on the outside.
Na+ Pump
- The Na+-K+ pump stabilizes the resting potential, because Na ions tend to lead inward and
K to leak outward along their concentration gradient
- The channel admits 2 potassium ions for every 3 sodium ions exported
- The movement of ions against the concentration gradient requires energy that is provided
by ATPase
Response to Stimulation: Action Potential
- Presynaptic neuron releases transmitter substance into the synaptic cleft by the arrival of
action potentials, or spikes
- The transmitter crosses the synaptic cleft and activate receptors of the target neuron
(postsynaptic) which activate transmitter-gated ion channel
- The activation of the ion channel alter the polarization of the target neuron
- If it is inhibitory it hyperpolarizes (by opening of Cl - channel), if it is excitatory it
depolarizes

Electrotonic Potentials
The initial target neuron response to excitatory stimulation is a local, graded or
electrotonic potential
At a low frequency of stimulation, small, decremental waves of depolarization extend for
50-100m along the affected dendrites, dying away after 2-3ms

With increasing frequency, the waves undergo temporal summation to form progressively
larger waves continuing on over the surface of the soma, spatial summation occurs when
waves traveling along two or more dendrites coalesce on the soma
About 15mV of depolarization, to a value of -55mV, brings the neuron to threshold (firing
level) at its most sensitive region, or trigger point, in the initial segment of the axon
The initial segment is the first region to give way at threshold voltage, because it is
exceptionally rich in voltage-gated sodium channels
When the level of depolarization (the generator potential) reaches threshold, nerve
impulses in the form of action potentials are suddently fired off
In sensory neurons of cranial and spinal nerve, the trigger zone generates what is known
as the receptor potential
The trigger zone of sensory neuron is exceptionally rich in the sensation-specific
transduction channel
In the case of myelinated nerve fibers, the trigger point is easily identified: in multipolar
neuron, it is immediately proximal to the first myelinated nerve fibers, and in peripheral
sensory neurons it is immediately distal to the final segment
The Shape of Action Potentials
When the local response to stimulation has depolarized the membrane to threshold, the
sudden increase in depolarization is brought about by the opening of voltage-gated sodium
channels
Sodium entry produces further depolarization and the positive feedback causes the
remaining sodium channels of the trigger zone to open, driving the membrane charge to
momentarily into a charge reversal (overshoot)
At this point, the sodium channels commence a progressive inactivation, and the voltagegated potassium channels are simultaneously triggered to open
Current flow switches from Na+ inflow to K+ outflow
The hyperpolarization phase (undershoot) is explained by the voltage-gated sodium
channels being completely inactivated prior to closure of the K + channels
Any remaining inconsistency is adjusted by activity of the Na +-K+ pump
In the resting state, an activation gate in the midregion of both Na + and K+ channel pores
is closed
The Na+ channel is the first to respond at threshold, by opening its activation gate and
allowing a torrential inflow of Na+ ions down the concentration gradient
One millisecond later, a second, inactivation gate, in the form of a flap of globular protein,
seals the exit into the cytosol while the K+ channel pore is opening
When the membrane potential approaches normality, the sodium gating reverts to its
resting inactive state
The action potential response to depolarization is all or none
It is different from graded potential is that it does not summate and it is not decremental
During the rising and early falling phases of the action potential, the neuron passes
through an absolute refractory period where it is incapable of initiating a second impulse
because too many voltage-gated channels are already open
This is followed by a relative refractory period where stimuli in excess of the standard
15mV requirement can elicit a response
It is quite common for the generator potential to reach up to 35mV, triggering impulses at
50-100 impulses per second, expressed as 50-100 Hz
Propagation
Depolarization at the trigger zone is propagated (conducted) along the axon
The membrane immediately proximal is sufficiently refractory to resist depolarization,
whereas that immediately distal undergoes a local response (depolarization) progressing
to firing level
This process continues distally along the stem axon and its branches, thereby conducting
the action potential all the way to the nerve terminals

Whereas conduction along unmyelinated nerve fibers is continuous, along myelinated

fibers it is salutatory (jumping)
Myelin sheaths are effective insulators overlying the intermodal segments, whereas Na
channels are very abundant at the nodes
Accordingly, spike potentials are generated at each successive node, the positive current
traveling along the axoplasm of the internode before exiting at the nex t node
As the current travels back through the ECF to recharge the depolarized patch of
membrane, withdrawal of positive charge causes the next node to depolarize