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Hyperpolarization (biology)
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Hyperpolarization is a change in a cell's membrane potential that makes it more

negative. It is the opposite of a depolarization. It inhibits action potentials by increasing
the stimulus required to move the membrane potential to the action potential threshold.
Hyperpolarization is often caused by efflux of K+ (a cation) through K+ channels, or influx
of Cl– (an anion) through Cl– channels. On the other hand, influx of cations,
e.g. Na+ through Na+ channels or Ca2+ through Ca2+ channels, inhibits hyperpolarization. If a
cell has Na+ or Ca2+ currents at rest, then inhibition of those currents will also result in a
hyperpolarization. This voltage-gated ion channel response is how the hyperpolarization
state is achieved. In neurons, the cell enters a state of hyperpolarization immediately
following the generation of an action potential. While hyperpolarized, the neuron is in a
refractory period that lasts roughly 2 milliseconds, during which the neuron is unable to
generate subsequent action potentials. Sodium-potassium ATPases redistribute K+ and
Na+ ions until the membrane potential is back to its resting potential of around –70
millivolts, at which point the neuron is once again ready to transmit another action
potential.[1]

Contents

 1Voltage-gated ion channels and hyperpolarization

 2Experimental technique
 3Examples
 4References
 5Further reading
Voltage-gated ion channels and hyperpolarization[edit]

The (a) resting membrane potential is a result of different concentrations of Na + and K+ ions inside and outside
the cell. A nerve impulse causes Na+ to enter the cell, resulting in (b) depolarization. At the peak action
potential, K+ channels open and the cell becomes (c) hyperpolarized.

Voltage gated ion channels respond to changes in the membrane potential. Voltage

gated potassium, chloride and sodium channels are key components in the generation
of the action potential as well as hyper-polarization. These channels work by selecting
an ion based on electrostatic attraction or repulsion allowing the ion to bind to the
channel.[2] This releases the water molecule attached to the channel and the ion is
passed through the pore. Voltage gated sodium channels open in response to a
stimulus and close again. This means the channel either is open or not, there is no part
way open. Sometimes the channel closes but is able to be reopened right away, known
as channel gating, or it can be closed without being able to be reopened right away,
known as channel inactivation.
At resting potential, both the voltage gated sodium and potassium channels are closed
but as the cell membrane becomes depolarized the voltage gated sodium channels
begin to open up and the neuron begins to depolarize, creating a current feedback
loop known as the Hodgkin cycle.[2] However, potassium ions naturally move out of the
cell and if the original depolarization event was not significant enough then the neuron
does not generate an action potential. If all the sodium channels are open, however,
then the neuron becomes ten times more permeable to sodium than potassium, quickly
depolarizing the cell to a peak of +40 mV.[2] At this level the sodium channels begin to
inactivate and voltage gated potassium channels begin to open. This combination of
closed sodium channels and open potassium channels leads to the neuron re-polarizing
and becoming negative again. The neuron continues to re-polarize until the cell reaches
~ –75 mV,[2] which is the equilibrium potential of potassium ions. This is the point at
which the neuron is hyperpolarized, between –70 mV and –75 mV. After
hyperpolarization the potassium channels close and the natural permeability of the
neuron to sodium and potassium allows the neuron to return to its resting potential of –
70 mV. During the refractory period, which is after hyper-polarization but before the
neuron has returned to its resting potential the neuron is capable of triggering an action
potential due to the sodium channels ability to be opened, however, because the neuron
is more negative it becomes more difficult to reach the action potential threshold.
HCN channels are activated by hyperpolarization.

Experimental technique[edit]

This image shows a model of a patch clamp used in neuroscience. The pipette tip is placed at an ion channel
opening and a current is applied and measured using a voltage clamp.

Hyperpolarization is a change in membrane potential, neuroscientists measure it using

a technique known as patch clamping. Using this method they are able to record ion
currents passing through individual channels. This is done using a glass micropipette,
also called a patch pipette, with a 1 micrometer diameter. There is a small patch that
contains a few ion channels and the rest is sealed off, making this the point of entry for
the current. Using an amplifier and a voltage clamp, which is an electronic feedback
circuit, allows the experimenter to maintain the membrane potential at a fixed point and
the voltage clamp then measures tiny changes in current flow. The membrane currents
giving rise to hyperpolarization are either an increase in outward current or a decrease
in inward current.[2]

Examples[edit]

Diagram of membrane potential changes during an action potential

 1. During the afterhyperpolarization period after
an action potential, the membrane potential is
more negative than when the cell is at the resting
potential. In the figure to the right, this
undershoot occurs at approximately 3 to 4
milliseconds (ms) on the time scale. The
afterhyperpolarization is the time when the
membrane potential is hyperpolarized relative to
the resting potential.
2. During the rising phase of an action potential, the
membrane potential changes from negative to
positive, a depolarization. In the figure, the rising
phase is from approximately 1 to 2 ms on the
graph. During the rising phase, once the
membrane potential becomes positive, the
membrane potential continues to depolarize
(overshoot) until the peak of the action potential
is reached at about +40 millivolts (mV). After the
peak of the action potential, a hyperpolarization
repolarizes the membrane potential to its resting
value, first by making it less positive, until 0 mV
is reached, and then by continuing to make it
more negative. This repolarization occurs in the
figure from approximately 2 to 3 ms on the time
scale.

References[edit]
1. ^ Pack, Phillip E. "Cliffs AP Biology 3rd Edition"
2. ^ Jump up to:          Becker, W. M., Kleinsmith, L. J., Hardin, J., &
a b c d e

Bertoni, G. P. (2009). Signal Transduction Mechanisms: I.

Electrical and Synaptic Signaling in Neurons. The World of
the Cell (7th ed., ). San Francisco: Pearson/Benjamin
Cummings.

Further reading[edit]
 Purves D, Augustine GJ, Fitzpatrick D, et al., eds.
(2001). Neuroscience (2. ed.). Sunderland, Mass:
Sinauer Assoc. ISBN 0-87893-742-0.
 Basic Neurochemistry Molecular, Cellular, and
Medical Aspects by Siegel, et al.
Categories: 
 Membrane biology
 Electrophysiology
  Electrochemistry
 Cellular neuroscience
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