inistry of higher education and M
cientific research s
l-Nahrain University A
ollege of medicine C
Electrical potentials
of nerves
By:
إبراهيم ماهر سلوم حسين
المرحلة األولى
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Introduction
Because electrical signals are the basis of
information transfer in the nervous system, it is
essential to understand how these signals arise. The
use of electrical signals—as in sending electricity
over wires to provide power or information—
presents a series of problems in electrical
engineering. A fundamental problem for neurons is
that their axons, which can be quite long (spinal
motor neuron can extend for a meter or more), are
not good electrical conductors. Although neurons
and wires are both capable of passively conducting
electricity, the electrical properties of neurons
compare poorly to even the most ordinary wire. To
compensate for this deficiency, neurons have
evolved a “booster system” that allows them to
conduct electrical signals over great distances
despite their intrinsically poor electrical
characteristics. The electrical signals produced by
this booster system are called action potentials
(which are also referred to as “spikes” or
.“impulses”)
The best way to observe an action potential is to use
an intracellular microelectrode to record directly the
electrical potential across the neuronal plasma membrane (As in figure). A typical
microelectrode is a piece of glass tubing pulled to a very fine point (with an
opening of 0.5 μm diameter) and filled with a good electrical conductor, such as a
concentrated salt solution. This conductive core can then be connected to a
voltmeter, such as an oscilloscope, to record the transmembrane voltage of the
nerve cell. When a microelectrode is inserted through the membrane of the neuron,
it records a negative potential, indicating that the cell has a means of generating a
constant voltage across its membrane when it is at rest. This voltage, called the
resting membrane potential, depends on the type of neuron being examined, but it
is always a fraction of a volt typically (-40 to -90 mV).
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This steady difference, the resting potential, is maintained whenever a neuron is
not generating impulses. The resting potential of a typical neuron is about -65
millivolts (1 mV = 0.001 volts). Stated another way, for a neuron at rest, Vm = 65
mV. This negative resting membrane potential of the neuron is an absolute
requirement for a functioning nervous system.
The Distribution of Ions Across the Membrane
The neuronal membrane potential depends on the ionic concentrations on both
sides of the membrane. Approximate values for these concentrations are shown
below. The important point is that K+ is more concentrated on the inside, and Na+
and Ca2+ are more concentrated on the outside. How do these concentration
gradients arise? Ionic concentration gradients are established by the actions of ion
pumps in the neuronal membrane. Two ion pumps are especially important in
cellular neurophysiology: the sodium-potassium pump and the calcium pump.
The sodium-potassium pump is an enzyme that breaks down ATP in the presence
of internal Na+. The chemical energy released by this reaction drives the pump,
which exchanges internal N+ for external K+. The actions of this pump ensure that
K+ is concentrated inside the neuron and that Na+ is concentrated outside. Notice
that the pump pushes these ions across the membrane against their concentration
gradients. This work requires the expenditure of metabolic energy. Indeed, it has
been estimated that the sodium-potassium pump expends as much as 70% of the
total amount of ATP utilized by the brain. The calcium pump is also an enzyme
that actively transports Ca+ out of the cytosol across the cell membrane. Additional
mechanisms decrease intracellular [Ca+] to a very low level (0.0002 mM); these
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include intracellular calcium-binding proteins and organelles, such as mitochondria
.and types of endoplasmic reticulum, which sequester cytosolic calcium ions
Ion pumps are the unsung heroes of cellular neurophysiology. They work in the
background to ensure that the ionic concentration gradients are established and
maintained. These proteins may lack the glamour of a gated ion channel, but
without ion pumps, the resting membrane potential would not exist and the brain
would not function.
Relative Ion Permeabilities of the Membrane at Rest
The pumps establish ionic concentration gradients across the neuronal membrane.
With knowledge of these ionic concentrations, we can use the Nernst equation to
calculate equilibrium potentials for the different ions.
RT [ion]
o
Nernst equation Eion =2.303 zF log [ion]
i
Where:
E ion: Ionic equilibrium potential
R: Gas constant
T: Absolute temperature
z: The charge of the ion
F: Faraday’s constant
Page | 4 [ion]o: Concentration of ion outside the cell
[ion]i: Concentration of ion inside the cell
The equilibrium potential for an ion is the membrane potential that would result if
a membrane were selectively permeable to that ion alone. In reality, however,
neurons are not permeable to only a single type of ion. How does that affect our
understanding?
Let’s consider a few scenarios involving K+ and Na+. If the membrane of a neuron
were permeable only to K+, the membrane potential would equal E K, which,
according to the table is 80 mV. On the other hand, if the membrane of a neuron
were permeable only to Na+, the membrane potential would equal E Na, 62 mV. If
the membrane were equally permeable to K+ and Na+, however, the resulting
membrane potential would be some average of E Na and E K. What if the membrane
were 40 times more permeable to K than it is to Na ? The membrane potential
again would be between E Na and E K but much closer to E K than to E Na. This
approximates the situation in real neurons. The actual resting membrane potential
of - 65 mV approaches, but does not reach, the potassium equilibrium potential of -
80 mV. This difference arises because, although the membrane at rest is highly
permeable to K+, there is also a steady leak of Na+ into the cell.
The resting membrane potential can be calculated using the Goldman equation, a
mathematical formula that takes into consideration the relative permeability of the
membrane to different ions. If we concern ourselves only with K+ and Na+, use the
ionic concentrations in the table, and assume that the resting membrane
permeability to K+ is fortyfold greater than it is to Na+, then the Goldman equation
predicts a resting membrane potential of - 65 mV, the observed value.
Where:
V: is the resulting potential
PNa, PK: is the relative permeabilities to Na and K respectively
The other terms are similar to Nernst equation
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Reference:
Neuroscience: Exploring the Brain 4th edition by Bear
PhD, Mark F., Connors PhD, Barry W., Paradiso
PhD (110-117).
Neuroscience, 2nd edition by Dale Purves, George J
Augustine, David Fitzpatrick, Lawrence C Katz,
Anthony-Samuel LaMantia, James O McNamara, and
S Mark Williams.
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