Resting membrane potential
The cell membrane of a neuron at rest has a
negative electrical potential of about -70mV. This means that if one were to
insert a voltmeter probe inside the cell, the electrical charges found there
and the charges found outside the cell would differ by 70mV, and the inside
would be negative relative to the outside. This electrical potential difference
is known as the resting membrane
potential(RMP). It is caused by a separation of charges across the membrane
differ, the membrane is said to be polarized.
The neuron has a high concentration of potassium ions(K+) on the
inside of the membrane and a high concentration of sodium ions(Na+)
on the outside. The imbalance in the number of ions inside and outside the cell
causes the RMP. The imbalance in the number of ions inside and outside the cell
causes the RMP. This imbalance is maintained in two ways. First, the cell membrane
is much more permeable to K+ than to Na+, so the K+
can move more freely. Because ions tend to move to establish equilibrium, some
of the K+ will move to an area where it is less concentrated,
outside the cell. The Na+ cannot move to the inside as easily.
Second, sodium-potassium pumps in
the neuron membrane, which contain Na+-K+ adenosine
triphosphatase(ATPase), maintain the imbalance on each side of the membrane by
actively transporting potassium ions
in the sodium ions out. The sodium-potassium pump moves three Na+
out of the cell for each two K+ it brings in. The end result is that
more positively charged ions are outside the cell than inside, creating the
potential difference across the membrane. Maintenance of a constant RMP of
about -70mV is primarily a function of the sodium-potassium pump.
Depolarization and hyperpolarization
If the inside of the cell becomes less negative
relative to the outside, the potential difference across the membrane
decreases. The membrane will be less polarized. When this happens, the membrane
is said to be depolarized. Thus, depolarization
occurs any time the charge difference becomes less than the RMP of -70mV,
moving closer to zero. This typically results from a change in the membrane’s
Na+ permeability.
The opposite can also occur. If the charge
difference across the membrane increases, moving from the RMP to an even more
negative value, then the membrane becomes more polarized. This is known as hyperpolarization. Changes in the
membrane potential are actually signals used to receive, transmit, and
integrate information within and between cells. These signals are of two types,
graded potentials and action
potentials. Both are electrical currents created by the movement of
ions.
Graded potentials
Graded
potentials are localized
changes in the membrane potential. These changes can be either depolarizations
or hyperpolarizations. The membrane contains ion channels with ion gates that
act as doorways into and out of the neuron. These gates are usually closed, preventing
ion flow; but they open with stimulation; allowing ions to move from the
outside to the inside or vice versa. This ion flow alters the charge
separation, changing the polarization of the membrane.
Graded potentials are triggered by a change in
the neuron’s local environment. Depending on the location and type of neuron
involved, the ion gates may open in response to the transmission of an impulse
from another neuron or in response to sensory stimuli such as changes in
chemical concentrations, temperature, or pressure.
Recall that most neuron receptors are located
on the dendrites( although some are on the cell body), yet the impulse is
always transmitted from the axon terminals at the opposite end of the cell. For
a neuron to transmit an impulse, the impulse must travel almost the entire
length of the neuron. Although a graded potential may result in depolarization
of the entire cell membrane, it is usually just a local event such that the
depolarization does not spread very far along the neuron. To travel the full
distance, an impulse must generate an action potential.
Action potentials
An action
potential is a rapid and substantial depolarization of the neuron’s
membrane. It usually lasts about 1ms. Typically, the membrane potential changes
from the RMP of about -70mV to a value of about +30mV and then rapidly returns
to its resting value. This is illustrated in the picture below. How does this
marked change in membrane potential occur?
All action potentials begin as graded
potentials. When enough stimulation occurs to cause a depolarization of at
least 15 to 20mV, an action potential results. In other words, if the membrane
depolarizes from the RMP of -70mV to a value of -50 to -55mV, the cell will
experience an action potential. The membrane voltage at which a graded
potential becomes an action potential is called the depolarization threshold.
Any depolarization that does not attain the threshold will not result in an
action potential. For example, if the membrane potential changes from the RMP of
-70mV to -60mV, the change is only 10mV and does not reach the threshold; thus,
no action potential occurs. But any time depolarization reaches or exceeds the
threshold, an action potential will result. This is the all-or-none principle.
When a given segment of an axon is generating
an action potential and its sodium gates are open, it is unable to respond to
another stimulus. This is reffered to as the absolute refractory period. When the sodium gates are
closed, the potassium gates are open, and repolarizing is occurring, the
segment of the axon can then respond to a new stimulus, but the stimulus must
be of substantially greater magnitude to evoke an action potential. This is
reffered to as the relative refractory
period.
Propagation of the action potential
Now that we understand how a neural impulse, in
the form of an action potential, is generated, we can look at how the impulse
is propagated, or how it travels through the neuron. Two characteristics of the
neuron become particularly important when we consider how quickly an impulse
can pass through the axon: myelination
and diameter.
Myelin
sheath
The axons of many neurons, especially large
neurons, are myelinated, meaning they are covered with a sheath formed by
myelin, a fatty substance that insulates the cell membrane. This myelin sheath is formed by specialized
cells called Schwann cells.
The sheath is not continuous. As it spans the
length of the axon, the myelin sheath exhibits gaps between adjacent Schwann
cells, leaving the axon uninsulated at those points. These gaps are reffered to
as nodes of Ranvier. The action
potential appears to jump from one node to the next as it transverses a
myelinated fiber. This is reffered to as salutatory
conduction, a much faster type of conduction than occurs in unmyelinated
fibers.
Myelination of peripheral motor neurons occurs
over the first several years of life, partly explaning why children need time
to develop coordinated movement. Individuals affected by certain neurological
diseases, such as MS as discussed in our chapter opening, experience
degeneration of the myelin sheath and a subsequent loss of coordination.
Diameter
of the neuron
The velocity of nerve impulse transmission is
also determined by the neuron’s size. Neurons of larger diameter conduct nerve
impulses faster than neurons of smaller diameter because larger neurons present
less resistance to local current flow.
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