Membrane Potentials

The action potential

10/30/2015
The main physiological effect of the electrical fields = electrical
stimulation of excitable tissue.
There are two types of electric response of the membrane to the
action of stimulus:
• the electrotonic (local) potential, in response to a weak
stimulus, below a certain threshold (subliminal)

• the action potential, which is a characteristic of excitable

cells, such as the neuron, the muscular fiber, the thrombocyte,
the sensory cells, and so on
Facts (related to the action potential):
• is an all-or-nothing response
• the threshold varies from one cell to another
• the initial and final values of the electrical potential are the
resting potential!

The action potential has 5 phases (!):

1. the resting potential (see Nernst/Goldman equations) (!)

• the equilibrium membrane potential – the net flow of all ions
across the membrane is zero!
• few potassium channels are open
• check permeability!
• there is NO NET movement of K ions (or any other ions)
2. threshold

• as soon as the depolarizing stimulus appears, membrane

potential becomes less negative
• both the voltage-gated sodium and potassium channels begin
to open!
◦ depolarization vs repolarization
▪ action potential vs electrotonic response
▪ all-or-none vs failed initiations
• strong stimuli vs weak stimuli

• when the threshold (RP + 15...25mV) is achieved, the action

potential is generated
3. the rising phase (typical 1ms)
• the depolarization reaches the threshold potential
• additional voltage-gated sodium channels are opened (positive
feedback) [V→ P → V]
• a high flow of Na ions enter into cell
• the membrane voltage becomes positive (U Na?)

4. the falling phase

• at the peak of the action potential, two processes occur
simultaneously:
◦ many voltage-gated sodium channels begin to close
▪ positive feedback → negative feedback (why?) - shut-off feature!!!
◦ many more potassium channels open / become fully open
• the positive charge leave the cell → the membrane potential
begins to shift back to the resting potential
• repolarization is speeded by voltage gated K channels
• as the membrane potential approaches the resting potential all
potassium channels (they react slow!) are open and maximally
activated → hyperpolarization / undershoot

5. the recovery phase

• the membrane repolarizes below the resting potential because
of number of the opened potassium channels (bigger than in
the resting state case)
• some (the additional ones) potassium channels close -
membrane’s permeability to potassium ions returns to its usual
value
Na+/K+ pump does not restore the resting potential!!!
After action potential - many sodium and potassium channels are
in refractory state (unable to open )!

Each action potential is followed by a refractory period:

• absolute refractory period (0.5...2ms) - it is impossible to
evoke another action potential
• relative refractory period - a stronger-than-usual stimulus is
required

• the refractory period...

◦ information travels only in one direction
◦ limits de maximum frequency of generating action
potentials
Facts:
• Na channels
◦ are fast
◦ voltage gated
◦ automatic inactivation – slow recovery
◦ 3 states: open, closed, inactivated
• K channels
◦ slower than Na channels
◦ voltage gated and simple
◦ no automatic inactivation
◦ 2 states: open, closed
 Long stimuli

Lapicque: How much and how long a stimulation is required to

excite a nerve?
strength – duration relation?
Lapicque’s law … Weiss equation

Is = a + b / t
(empirical equation)

Is = IR / ( 1 – e-t/T )
IR = rheobase
TM = RM CM = the membrane time
constant (~1ms)
2IR → chronaxie
IR depends on
• physiological parameters
• individual exposure conditions

chronaxie = 0.01ms (motoneuron)

chronaxie = 0.35ms (sensory neuron)
chronaxie = 0.8ms (skin)
chronaxie = 100ms (smooth muscle - frog)

Weiss-Lapicque’s law:
• Stimuli must exceed a threshold in order to create an action potential
• Stimuli below the threshold cannot create an action potential (even if they are of
very long duration)
• Stimuli with shorter durations must be of higher intensity in order to be effective