Chapter 7 PDF

Chapter 7 https://droualb.faculty.mjc.edu/Course Materials/Physiology 10...
Chapter 7 - Nerve Cells and Electric Signaling
Overview
Organization:
Central Nervous System (CNS)
Brain and spinal cord receives and processes information.
Peripheral Nervous System (PNS)
Nerve cells that link CNS with organs throughout the body. Neurons that transmit messages to an
organ or carry information from a sensory organ are said to innervate that organ.
Divisions:
a. Afferent
Provides information about the somatic senses (the body in general as we are aware of it)
including information from the skin, muscles and joints; special senses (vision, hearing,
equilibrium, taste and smell), and visceral senses (information from the internal organs).
b. Efferent
Sends out commands to effector organs (muscles and glands).
Subdivisions:
1. Somatic Nervous System
Include nerve cells that innervate skeletal muscle.
2. Autonomic Nervous System
Nerve cells that control smooth muscle and glands which are not under voluntary control.
Enteric Nervous System

Network of nerves in gastrointestinal tract that can function independently of the rest of the nervous
system but which is under the influence of the autonomic nervous system.
Cells of the Nervous System
Two kinds of cells:
1. Neuron or nerve cell is the functional unit. These cells are excitable cells capable of conducting
electrochemical signals along their membranes.
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2. Glial cells provide structural and metabolic support for neurons. These cells constitute about 90%
of the cells in the nervous system.
Neurons
Consist of:
Cell body (soma) - Contains the nucleus and most of the cell's organelles.
Dendrites - Processes that branch off the cell body and receive input from other neurons at specialized
junctions called synapses. The cell body also receives this input.
Axons (nerve fiber) - Processes that send information. Typically each neuron has only one axon
coming off the cell body but the axon may have branches called collaterals.
Axons transmit information over long distances in the form of action potentials. The axon comes off
the cell body at the axon hillock and transmits an action potential to the axon terminal where the
neuron synapses with another neuron or effector organ.
Locations of Ion Channels

Leak channels are non-gated channels found throughout the neuron. They are always open and
contribute to the resting membrane potential.
Ligand-gated channels are found on dendrites and the cell body and open or close in response to
the presence of neurotransmitters (ligands).
Voltage-gated channels open or close in response to changes in membrane potential. Many of these
are voltage-gated Na+ and K+ channels that are important for the initiation and propagation of action
potentials along the axon. Voltage-gated Ca2+ channels are found at axon terminals where the opening
of these channels permit Ca2+ to enter and trigger the release of neurotransmitter.
Classification of Neurons
Structural Classification
Bipolar
A neuron that has two processes, one axon and one dendrite. Seen in special sensory neurons for
olfaction and vision.
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Most sensory neurons are a subclass of bipolar neurons called pseudo-unipolar. This is a neuron
with one process coming off the cell body. However, this process results from the fusion of an axon and
a dendrite modified to transmit action potentials.
Multipolar
This neuron is the most common. The multipolar neuron has multiple processes with one being an
axon and the rest dendrites.
Functional Classification
Afferent Neuron
Neurons that transmit sensory information from the outside (sensory receptors) or inside (visceral
receptors) the body to the CNS.
Interneurons
Neurons located entirely in the CNS that process sensory information; send out commands to effector
organs; and perform complex integrative and analytical functions. Most neurons (99%) of the nervous
system are interneurons.
Efferent Neuron
Neurons that transmit information or commands from the CNS to effector organs.
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Glial Cells
Glial cells include astrocytes, ependymal cells, microglia, oligodendrocytes (in the CNS) and Schwann
cells (in the PNS). Oligodendrocytes and Schwann cells increase the rapidity and efficiency of nerve
transmission by forming an insulating wrap of myelin around the axons of neurons. Myelin consists of
concentric layers of cell membranes wrapped around axons. Myelin forms a barrier that prevents the leakage
of ions except at gaps between consecutive oligodendrocytes and Schwann cells called nodes of Ranvier.
The nodes of Ranvier contain voltage-gated sodium and potassium channels that function in the transmission
of action potentials.
Electrical Concepts
Electrical potentials (E) result from the separation of opposite charges (measured in millivolts). The
actual movement of charges is called current (measured in microamperes).
Resistance (R ) is a measure of hindrance to the movement of charged particles. Charge flow through
extra and intracellular fluid with ease but the lipid bilayer of cell membranes create a barrier to the
movement of charge.
Conductance is a measure of the ability of charges to move across a membrane and is inversely related
to resistance:
g = 1/R
Ohm's Law describes the relationship between potential difference, current and resistance and is
expressed in the equation
I = E/R
where I = current, E = potential difference and R = resistance.
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Resting Membrane Potential Animation showing creation of resting membrane potential

A nerve cell at rest (not undergoing an electrochemical change across its membrane) has a resting membrane
potential of approximately - 70 mV across its membrane. (This actually varies among different cells but unless
otherwise indicated we will assume this to be the value.) This is because across the cell membrane of the neuron the
intracellular surface of the membrane is -70 mV more negative than the extracellular surface.
The resting membrane potential results from the concentration gradients of ions across the cell membrane and
the presence of ion channels in the plasma membrane.
The Na+/K+ pump pumps 3 Na+ out of the cell for every 2 K+ it pumps in creating concentration gradients for
Na+ and K+ across the cell membrane. Na+ is more concentrated on the outside, and K+ is more concentrated on the
inside.
Also present in the cell membrane are ion channels for Na+ and K+ as well as for Cl- and Ca++. The resting
membrane potential depends on the permeability of the cell membrane for these ions, particularly Na+ and K+. The
permeability of these ions depends upon the presence of the ion channels and whether or not they are open.
Equilibrium potential for K+
If the cell were only permeable to K+, K+ would leave the cell and go down its concentration gradient. As the
positively charged K+ leaves the cell, a negative membrane potential develops. This results in two forces acting
upon K+ , the chemical force (concentration gradient) pushing K+ out, and an electrical force (negative
membrane potential) pulling the K+ in. The two forces together constitute an electrochemical force and balance
one another when the membrane potential is -94 mV. Hence, the equilibrium potential for K+ is -94 mV, or, Ek+
= -94 mV.
Equilibrium potential for Na+
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If the cell is only permeable to Na+ this ion will move across the membrane down its concentration gradient.
This creates a positive membrane potential that will balance the chemical force when the membrane potential is
+60 mV. Hence, ENa+ = 60 mV.
Resting Membrane Potential of Neurons The Nernst-Goldman Equation Simulator
The real cell has similar concentration gradients of Na+ and K+ across the membrane. However, when the
membrane is resting, K+ is about 25 times more permeable than Na+. Both K+ and Na+ will move down their
concentration gradients but in opposite directions. This movement of K+ out of the cell, and Na+ into the cell,
continues until the number of positive charges exiting the cell equals the number of positive charges entering. At
this point the resting membrane potential is reached at - 70 mV.
The final resting membrane potential reflects a balance between the electrochemical forces associated with each
ion. When the cell is "at rest", the cell is more permeable to K+ than to Na+ and because of this, the electrochemical
force of K+ has a greater influence over the membrane potential than the electrochemical force of Na+. The resting
potential of -70 mV is a lot closer to EK+ than to ENa+.
The resting membrane potential needs to be maintained by the Na+/K+ pump that is constantly pumping Na+ out
and K+ in. This is because over time Na+ would continually leak in and K+ would leak out and the concentration
gradients would diminish. The Na+/K+ pump keeps the membrane potential as a steady state. Because more Na+
(3) are pumped out than K+ (2) are pumped in, the Na+/K+ pump is electrogenic, but this makes only a small
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contribution to the final resting potential.

The general rule is that a cell's membrane potential is a weighted sum of the equilibrium potentials of all
permeant ions. The weighting (amount of influence) given to each ion is proportional to that ion's permeability.
The more permeable the ion, the closer the resting potential will match that of the ion.
Another important concept to understand is that at the steady state of the resting membrane potential, the ions are
not at their equilibrium potentials and there is an electrochemical force acting on the ions to bring them to
equilibrium. The net electrochemical force driving the ions is proportional to the difference between the membrane
potential and the equilibrium potential for that ion.
Hence, at a resting membrane potential of -70 mV K+ is -24 mV (-94 mV - (-70) mV = -24 mV) away from its
equilibrium potential and Na+ is 130 mV (60 mV - (- 70) mV = 130 mV) from its equilibrium potential. In other
words, the electrochemical force acting upon Na+ to move it into the cell is greater than that acting upon K+ to move
it out of the cell.
Changes in Membrane Potential
The resting membrane potential is due to the difference in the permeability of the cell membrane to Na+ and K+
as determined by leak channels for these respective ions. Changes in the membrane potential from the resting level
result from gated ion channels.
Gated ion channels are channels that open in response to some stimulus as indicated below:
1. Electrical changes - Voltage-gated channels.
2. Chemical messengers - Ligand-gated channels.
3. Physical alteration - Mechanically-gated channels
The typical nerve cell has a resting potential of -70 mV. At this resting level the membrane is polarized because
the inside surface of the membrane is negative with respect to the outside. If the potential becomes more negative
(e.g. -90 mV) the membrane is said to be hyperpolarized. When the membrane becomes less negative (e.g. -50
mV) the membrane is said to be depolarized. After the membrane is depolarized, when it returns to its resting level
it is repolarized.
Electrical Signaling Through Changes in the Membrane Potential Animation of Action Potential from Harvard
Neurons communicate through two kinds of changes in membrane potential:
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1. Graded Potentials
A graded potential is a small change in the potential that is proportional (graded) to the strength of the
stimulus causing the change. The stimulus may be a neurotransmitter or a sensory stimulus. Graded
potentials may be either depolarization or hyperpolarizations.
Graded potentials are significant because they either bring the membrane potential closer to or further from
the potential that triggers an action potential. The potential that triggers an action potential is the threshold. If
the graded potential brings the potential closer to threshold it is excitatory. If it brings it further from
threshold it is inhibitory.
A graded potential can only travel a short distance because the change in voltage spreads by the passive
movement of ions in a process called electrotonic conduction. As the current moves further from the site of
stimulation, the membrane potential decreases because the ions diffuse and the ions pass through channels in
the membrane. Therefore, the change in membrane potential that is due to electrotonic conduction is
decremental (decreasing).
The graded potentials produced in neurons may add up spatially or temporally:
In spatial summation the graded potentials produced at different locations of the cell membrane add
together. This addition is particularly effective if the graded potentials are produced about the same
time. It is important to remember that the graded potentials can be inhibitory (hyperpolarizing) as well
as excitatory (depolarizing) and can therefore cancel each other out.
In temporal summation the stimuli producing the graded potential at the same location come together
so rapidly the postsynaptic membrane does not have time to repolarize before the next stimulus is
received. The greater the overlap in time, the greater the temporal summation.
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2. Action Potentials
During an action potential a rapid depolarization occurs that actually causes a reversal in the polarity of
the membrane (-70 mV to 30 mV). This reversal is brief and the negative potential is rapidly restored. Once
the action potential begins it becomes self propagating and can travel over a long distance along the length of
an axon without any decrease in strength. (See table 7.2 for comparisons between graded and action
potentials.)
Ionic Basis Action Potential Animation
There are three distinct phases in an action potential that depends upon the electrochemical gradients of
Na and K+ and changes in the permeability of the membrane to these ions.
+
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1. Depolarization. A sudden increase in the permeability of the membrane to Na+ causes Na+ ions to
rush into the cell. The permeability of Na+ is now greater than that of K+ and the membrane potential
swings toward the equilibrium potential for Na+ (+60 mV). The membrane potential goes from -70 mV
to 30 mV.
2. Repolarization. Within 1 msec Na+ permeability decreases rapidly and the K+ permeability
increases. This causes a net outflow of positive charge as K+ moves down its electrochemical gradient
and the membrane potential becomes negative again returning to -70 mV.
3. After-Hyperpolarization. The potassium permeability remains high for 5-15 msec. This causes the
membrane to overshoot the resting membrane potential and hyperpolarize as the increase in K+
permeability causes the membrane potential to approach the equilibrium potential of K+ (-94 mV).
The changes in permeability that occur are due to voltage-gated ion channels.
Voltage-Gated Ion Channels
Voltage-gated Na+ and K+ channels are located primarily in the plasma membrane of the axon hillock and
axon. A model is used to explain how these channels work. The model for the Na+ channel states that there
are two kinds of gates in the Na+ channels:
1. Activation gate - opens to allow ions to pass through.
2. Inactivation gate - closes to block the passage of ions.
If we take the voltage-gated channel for Na+ the model will work this way during an action potential: Go
to: Voltage-gated channels and the action potential
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a. During the resting membrane potential the activation gate is closed while the inactivation gate is
open.
b. During the depolarization phase, the activation gate opens with both gates open Na+ rushes down
its concentration gradient and enters the cell.
c. When repolarization begins (approximately 1 msec after the activation gate opens) the inactivation
gate closes and stops the inrush of Na+.
d. When repolarization ends and the membrane returns to resting potential the activation gate closes
and the inactivation gate opens. The channel has now returned to its original position before
depolarization.
The activation of the voltage-gated Na+ channels is a regenerating phenomenon because when activated
the change in potential activates other channels in a positive feedback loop. This is because when one
channel opens and Na+ rushes in, this depolarizes the membrane and activates other Na+ channels. These
channels in turn activate more channels, and so forth.
The potassium channels contribute to the repolarizing phase of the action potential by opening slowly and
closing slowly. The model for the voltage-gated K+ channel describes only a single gate that opens slowly in
response to depolarization. As the K+ flows out and repolarizes the membrane these slowly close.
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The threshold for an action potential is reached when the opening voltage-gated sodium channels
stimulates other channels to open in a positive feedback loop. A depolarization that does not generate an
action potential is subthreshold. A depolarization that is above threshold elicits the same action potential as
one that just reaches threshold.
Action potentials are initiated according to an all-or-none principle. Either the depolarization does not
reach threshold and triggers no action potential or it reaches threshold and elicits an action potential that is
always the same.
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Refractory Periods
After an action potential there is a refractory period when the membrane is not as excitable. There are two
kinds of refractory periods:
1. Absolute Refractory Period occurs 1-2 msec after the initiation of the action potential. During this
time an action potential cannot be generated.
2. Relative Refractory Period immediately follows the absolute refractory period and lasts 5 - 15 msec.
During this time a second action potential can be generated but only if a stronger stimulus is used.
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The absolute refractory period contributes to the all-or-none property of action potentials by making it
impossible for action potentials to add-up like graded potentials. Relative refractory periods make it possible
to encode information by converting the strength of a stimulus into the frequency of action potentials. A
suprathreshold graded potential can produce a higher number of action potentials within a given period of time
then a lesser subthreshold or threshold stimulus.
A threshold graded potential can produce more than one action potential if it lasts for a longer period of time
than the refractory period. A suprathreshold graded potential produces more action potentials by stimulating the
membrane enough during the relative refractory period to elicit action potentials. The absolute refractory period
imposes a maximum frequency of action potentials of 500 - 1000 per second.
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Propagation of Action Potentials Propagation of the Action Potential;

Unmyelinated Axon Go to: Action potential propagation in an unmyelinated axon
In unmyelinated axons, action potentials are spread by electrotonic conduction or the passive spread of
voltage change along the membrane of the axon. When an action potential occurs at the trigger site (axon
hillock) positive charges rush into the cell. This creates a local zone in both the extracellular and intracellular
fluid where there is a sudden change in charge. If we concentrate only on the intracellular fluid, the positive
charges that enter the cell move toward the surrounding areas where there is a high concentration of negative
charges. This results in a depolarization that reaches threshold and continues the action potential.
The action potential moves along the axon in one direction because of the refractory period. The action
potential travels continuously along the length of the axon like a wave.
The larger the diameter of the unmyelinated neuron the less the resistance to current flow and the faster
the propagation of the action potential.
Myelinated Axon
Myelin provides high resistance to ion flow across the membrane. This resistance is only lacking where
there is a gap in this myelin covering at the node of Ranvier. Also at the nodes of Ranvier there is a
concentration of voltage-gated sodium and potassium channels.
When an action potential occurs at one node the same intracellular and extracellular currents are created
except that the myelin dramatically reduces the current across the membrane so that the current that flows to
the next node is strong enough to generate an action potential. This continues down the length of a
myelinated axon with the action potential jumping from one node to the next in what is called saltatory
conduction.
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Saltatory conduction enables the action potential to travel faster. Hence, the larger the axon diameter the
faster the conduction and myelination speeds up conduction speed even more. Conduction velocity is
associated with nerve function (Table 7.4).
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Chapter 7 https://droualb.faculty.mjc.edu/Course Materials/Physiology 10...

Chapter 7 - Nerve Cells and Electric Signaling

Enteric Nervous System

Locations of Ion Channels

where I = current, E = potential difference and R = resistance.

Resting Membrane Potential Animation showing creation of resting membrane potential

Equilibrium potential for K+

Resting Membrane Potential of Neurons The Nernst-Goldman Equation Simulator

contribution to the final resting potential.

Propagation of Action Potentials Propagation of the Action Potential;

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