2. Nerve and Muscle Cells
2Nerve and Muscle
Cells
2.1 INTRODUCTION
In this chapter we consider the structure of nerve and muscle
tissue and in particular their membranes, which are excitable. A qualitative
description of the activation process follows. Many new terms and concepts are
mentioned only briefly in this chapter but in more detail in the next two
chapters, where the same material is dealt with from a quantitative rather than
a qualitative point of view. The first documented
reference to the nervous system is found in ancient Egyptian records. The Edwin
Smith Surgical Papyrus, a copy (dated 1700 B.C.) of a manuscript composed about
3500 B.C., contains the first use of the word "brain", along with a description
of the coverings of the brain which was likened to the film and corrugations
that are seen on the surface of molten copper as it cooled (Elsberg, 1931;
Kandel and Schwartz, 1985). The basic unit of living
tissue is the cell. Cells are specialized in their anatomy and physiology to
perform different tasks. All cells exhibit a voltage difference across the cell
membrane. Nerve cells and muscle cells are excitable. Their cell membrane can
produce electrochemical impulses and conduct them along the membrane. In muscle
cells, this electric phenomenon is also associated with the contraction of the
cell. In other cells, such as gland cells and ciliated cells, it is believed
that the membrane voltage is important to the execution of cell function.
The origin of the
membrane voltage is the same in nerve cells as in muscle cells. In both cell
types, the membrane generates an impulse as a consequence of excitation. This
impulse propagates in both cell types in the same manner. What follows is a
short introduction to the anatomy and physiology of nerve cells. The reader can
find more detailed information about these questions in other sources such as
Berne and Levy (1988), Ganong (1991), Guyton (1992), Patton et al. (1989) and
Ruch and Patton (1982).
2.2 NERVE CELL
2.2.1 The Main Parts of the Nerve Cell
The nerve cell may be divided on the basis of its structure and
function into three main parts: (1) the cell body,
also called the soma; (2) numerous short processes of the soma, called the
dendrites; and, (3) the single long nerve fiber, the axon.
These are described in Figure 2.1. The body of a nerve cell
(see also (Schadé and Ford, 1973)) is similar to that of all other cells. The
cell body generally includes the nucleus, mitochondria, endoplasmic reticulum,
ribosomes, and other organelles. Since these are not unique to the nerve cell,
they are not discussed further here. Nerve cells are about 70 - 80% water; the
dry material is about 80% protein and 20% lipid. The cell volume varies between
600 and 70,000 µmÅ‚. (Schadé and Ford, 1973) The short processes of the
cell body, the dendrites, receive impulses from other cells and transfer them to
the cell body (afferent signals). The effect of these impulses may be
excitatory or inhibitory. A cortical neuron (shown in
Figure 2.2) may receive impulses from tens or even hundreds of thousands of
neurons (Nunez, 1981). The long nerve fiber, the axon, transfers the signal from the
cell body to another nerve or to a muscle cell. Mammalian axons are usually
about 1 - 20 µm in diameter. Some axons in larger animals may be several meters
in length. The axon may be covered with an insulating layer called the myelin
sheath, which is formed by Schwann cells (named for the German
physiologist Theodor Schwann, 1810-1882, who first observed the myelin sheath in
1838). The myelin sheath is not continuous but divided into sections, separated
at regular intervals by the nodes of Ranvier (named for the French
anatomist Louis Antoine Ranvier, 1834-1922, who observed them in 1878).
Fig. 2.1. The major components of a neuron.
Fig. 2.2. Cortical nerve cell and nerve endings connected to it.
2.2.2 The Cell Membrane
The cell is enclosed by a cell membrane whose thickness is
about 7.5 - 10.0 nm. Its structure and composition resemble a soap-bubble film
(Thompson, 1985), since one of its major constituents, fatty acids, has that
appearance. The fatty acids that constitute most of the cell membrane are called
phosphoglycerides. A phosphoglyceride consists of phosphoric acid and
fatty acids called glycerides (see Figure 2.3). The head of this
molecule, the phosphoglyceride, is hydrophilic (attracted to water). The
fatty acids have tails consisting of hydrocarbon chains which are
hydrophobic (repelled by water). If fatty acid molecules
are placed in water, they form little clumps, with the acid heads that are
attracted to water on the outside, and the hydrocarbon tails that are repelled
by water on the inside. If these molecules are very carefully placed on a water
surface, they orient themselves so that all acid heads are in the water and all
hydrocarbon tails protrude from it. If another layer of molecules were added and
more water put on top, the hydrocarbon tails would line up with those from the
first layer, to form a double (two molecules thick) layer. The acid heads would
protrude into the water on each side and the hydrocarbons would fill the space
between. This bilayer is the basic structure of the cell membrane. From the bioelectric
viewpoint, the ionic channels constitute an important part of the cell
membrane. These are macromolecular pores through which sodium, potassium, and
chloride ions flow through the membrane. The flow of these ions forms the basis
of bioelectric phenomena. Figure 2.4 illustrates the construction of a cell
membrane.
Fig. 2.3. A sketch illustrating how the phosphoglyceride (or
phospholipid) molecules behave in water. See text for discussion.
Fig. 2.4. The construction of a cell membrane. The main
constituents are two lipid layers, with the hydrophobic tails pointing inside
the membrane (away from the aqueous intracellular and interstitial mediums).
The macromolecular pores in the cell membrane form the ionic channels through
which sodium, potassium, and chloride molecules flow through the membrane and
generate the bioelectric phenomena.
2.2.3 The Synapse
The junction between an axon and the next cell with which it
communicates is called the synapse. Information proceeds from the cell
body unidirectionally over the synapse, first along the axon and then across the
synapse to the next nerve or muscle cell. The part of the synapse that is on the
side of the axon is called the presynaptic terminal; that part on the
side of the adjacent cell is called the postsynaptic terminal. Between
these terminals, there exists a gap, the synaptic cleft, with a thickness of 10
- 50 nm. The fact that the impulse transfers across the synapse only in one
direction, from the presynaptic terminal to the postsynaptic terminal, is due to
the release of a chemical transmitter by the presynaptic cell. This transmitter,
when released, activates the postsynaptic terminal, as shown in Figure 2.5. The
synapse between a motor nerve and the muscle it innervates is called the
neuromuscular junction. Information transfer in the synapse is discussed
in more detail in Chapter 5.
Fig. 2.5. Simplified illustration of the anatomy of the
synapse.(A) The synaptic vesicles contain a chemical transmitter.(B)
When the activation reaches the presynaptic terminal the transmitter is
released and it diffuses across the synaptic cleft to activate the
postsynaptic membrane.
2.3 MUSCLE CELL
There are three types of muscles in the body: - smooth muscle, - striated muscle
(skeletal muscle), and - cardiac muscle. Smooth muscles are involuntary (i.e., they cannot be
controlled voluntarily). Their cells have a variable length but are in the order
of 0.1 mm. Smooth muscles exist, for example, in the digestive tract, in the
wall of the trachea, uterus, and bladder. The contraction of smooth muscle is
controlled from the brain through the autonomic nervous system. Striated muscles,
are also called skeletal muscles because of their anatomical location,
are formed from a large number of muscle fibers, that range in length from 1 to
40 mm and in diameter from 0.01 to 0.1 mm. Each fiber forms a (muscle) cell and
is distinguished by the presence of alternating dark and light bands. This is
the origin of the description "striated," as an alternate terminology of
skeletal muscle (see Figure 2.6). The striated muscle fiber
corresponds to an (unmyelinated) nerve fiber but is distinguished
electrophysiologically from nerve by the presence of a periodic transverse
tubular system (TTS), a complex structure that, in effect, continues the surface
membrane into the interior of the muscle. Propagation of the impulse over the
surface membrane continues radially into the fiber via the TTS, and forms the
trigger of myofibrillar contraction. The presence of the TTS affects conduction
of the muscle fiber so that it differs (although only slightly) from propagation
on an (unmyelinated) nerve fiber. Striated muscles are connected to the bones
via tendons. Such muscles are voluntary and form an essential part of the organ
of support and motion. Cardiac muscle is also striated, but differs in other ways from
skeletal muscle: Not only is it involuntary, but also when excited, it generates
a much longer electric impulse than does skeletal muscle, lasting about 300 ms.
Correspondingly, the mechanical contraction also lasts longer. Furthermore,
cardiac muscle has a special property: The electric activity of one muscle cell
spreads to all other surrounding muscle cells, owing to an elaborate system of
intercellular junctions.
Fig. 2.6. Anatomy of striated muscle. The fundamental physiological
unit is the fiber.
2.4 BIOELECTRIC FUNCTION OF THE NERVE CELL
The membrane voltage (transmembrane voltage)
(Vm) of an excitable cell is defined as the potential at the
inner surface (Fi)
relative to that at the outer (Fo) surface of the membrane, i.e.
Vm = (Fi) - (Fo). This definition is independent of the
cause of the potential, and whether the membrane voltage is constant, periodic,
or nonperiodic in behavior. Fluctuations in the membrane potential may be
classified according to their character in many different ways. Figure 2.7 shows
the classification for nerve cells developed by Theodore Holmes Bullock (1959).
According to Bullock, these transmembrane potentials may be resolved into a
resting potential and potential changes due to activity. The latter may be
classified into three different types:
1. Pacemaker potentials: the intrinsic activity of the cell which
occurs without external excitation.
2. Transducer potentials across the membrane, due to
external events. These include generator potentials caused by receptors
or synaptic potential changes arising at synapses. Both subtypes can be
inhibitory or excitatory.
3. As a consequence of transducer potentials, further
response will arise. If the magnitude does not exceed the threshold, the
response will be nonpropagating (electrotonic). If the response is
great enough, a nerve impulse (action potential impulse) will be
produced which obeys the all-or-nothing law (see below) and proceeds
unattenuated along the axon or fiber.
Fig. 2.7. Transmembrane potentials according to Theodore H.
Bullock.
2.5 EXCITABILITY OF NERVE CELL
If a nerve cell is stimulated, the transmembrane voltage
necessarily changes. The stimulation may be
excitatory (i.e., depolarizing; characterized by a change of
the potential inside the cell relative to the outside in the positive
direction, and hence by a decrease in the normally negative resting voltage)
orinhibitory (i.e., hyperpolarizing, characterized by a
change in the potential inside the cell relative to the outside in the
negative direction, and hence by an increase in the magnitude of the membrane
voltage).
After stimulation the membrane voltage returns to its original
resting value. If
the membrane stimulus is insufficient to cause the transmembrane potential to
reach the threshold, then the membrane will not activate. The response of the
membrane to this kind of stimulus is essentially passive. Notable research on
membrane behavior under subthreshold conditions has been performed by
Lorente de Nó (1947) and Davis and Lorente de Nó (1947). If the excitatory stimulus
is strong enough, the transmembrane potential reaches the threshold, and the
membrane produces a characteristic electric impulse, the nerve impulse.
This potential response follows a characteristic form regardless of the strength
of the transthreshold stimulus. It is said that the action impulse of an
activated membrane follows an all-or-nothing law. An inhibitory stimulus
increases the amount of concurrent excitatory stimulus necessary for achieving
the threshold (see Figure 2.8). (The electric recording of the nerve impulse is
called the action potential. If the nerve impulse is recorded
magnetically, it may be called an action current. The terminology is
further explicated in Section 2.8 and in Figure 2.11, below.)
Fig. 2.8. (A) Experimental arrangement for measuring the response
of the membrane potential (B) to inhibitory (1) and excitatory (2, 3, 4)
stimuli (C). The current stimulus (2), while excitatory is, however,
subthreshold, and only a passive response is seen. For the excitatory level
(3), threshold is marginally reached; the membrane is sometimes activated
(3b), whereas at other times only a local response (3a) is seen. For a
stimulus (4), which is clearly transthreshold, a nerve impulse is invariably
initiated.
2.6 THE GENERATION OF THE ACTIVATION
The mechanism of the activation is discussed in detail in
Chapter 4 in connection with the Hodgkin-Huxley membrane model. Here the
generation of the activation is discussed only in general terms. The concentration of
sodium ions (Na+) is about 10 times higher outside the membrane than
inside, whereas the concentration of the potassium (K+) ions is about
30 times higher inside as compared to outside. When the membrane is stimulated
so that the transmembrane potential rises about 20 mV and reaches the threshold
- that is, when the membrane voltage changes from -70 mV to about -50 mV (these
are illustrative and common numerical values) - the sodium and potassium ionic
permeabilities of the membrane change. The sodium ion permeability increases
very rapidly at first, allowing sodium ions to flow from outside to inside,
making the inside more positive. The inside reaches a potential of about +20 mV.
After that, the more slowly increasing potassium ion permeability allows
potassium ions to flow from inside to outside, thus returning the intracellular
potential to its resting value. The maximum excursion of the membrane voltage
during activation is about 100 mV; the duration of the nerve impulse is around 1
ms, as illustrated in Figure 2.9. While at rest, following activation, the Na-K
pump restores the ion concentrations inside and outside the membrane to their
original values.
Fig. 2.9. Nerve impulse recorded from a cat motoneuron following a
transthreshold stimulus. The stimulus artifact may be seen at t = 0.
2.7 CONCEPTS ASSOCIATED WITH THE ACTIVATION
PROCESS
Some basic concepts associated with the activation process are
briefly defined in this section. Whether an excitatory cell is activated depends
largely on the strength and duration of the stimulus. The membrane potential may
reach the threshold by a short, strong stimulus or a longer, weaker stimulus.
The curve illustrating this dependence is called the strength-duration
curve; a typical relationship between these variables is illustrated in
Figure 2.10. The smallest current adequate to initiate activation is called the
rheobasic current or rheobase. Theoretically, the rheobasic
current needs an infinite duration to trigger activation. The time needed to
excite the cell with twice rheobase current is called chronaxy. Accommodation and
habituation denote the adaptation of the cell to a continuing or
repetitive stimulus. This is characterized by a rise in the excitation
threshold. Facilitation denotes an increase in the excitability of the
cell; correspondingly, there is a decrease in the threshold. Latency
denotes the delay between two events. In the present context, it refers to the
time between application of a stimulus pulse and the beginning of the
activation. Once activation has been initiated, the membrane is insensitive to
new stimuli, no matter how large the magnitude. This phase is called the
absolute refractory period. Near the end of the activation impulse, the
cell may be activated, but only with a stimulus stronger than normal. This phase
is called the relative refractory period. The activation process
encompasses certain specifics such as currents, potentials, conductivities,
concentrations, ion flows, and so on. The term action impulse describes
the whole process. When activation occurs in a nerve cell, it is called a
nerve impulse; correspondingly, in a muscle cell, it is called a
muscle impulse. The bioelectric measurements focus on the
electric potential difference across the membrane; thus the electric
measurement of the action impulse is called the action potential that
describes the behavior of the membrane potential during the activation.
Consequently, we speak, for instance, of excitatory postsynaptic
potentials (EPSP) and inhibitory postsynaptic potentials (IPSP). In
biomagnetic measurements, it is the electric current that is the
source of the magnetic field. Therefore, it is logical to use the term action
current to refer to the source of the biomagnetic signal during the action
impulse. These terms are further illustrated in Figure 2.11.
Fig. 2.10. (A) The response of the membrane to various stimuli of
changing strength (B), the strength-duration curve. The level of current
strength which will just elicit activation after a very long stimulus is
called rheobase. The minimum time required for a stimulus pulse twice the
rheobase in strength to trigger activation is called chronaxy. (For
simplicity, here, threshold is shown to be independent on stimulus duration.)
Fig. 2.11. Clarification of the terminology used in connection with
the action impulse: A) The source of the action impulse may be nerve or
muscle cell. Correspondingly it is called a nerve impulse or a muscle impulse.
B) The electric quantity measured from the action impulse may be potential
or current. Correspondingly the recording is called an action potential or an
action current.
2.8 CONDUCTION OF THE NERVE IMPULSE IN AN
AXON
Ludvig Hermann (1872, 1905) correctly proposed that the
activation propagates in an axon as an unattenuated nerve impulse. He suggested
that the potential difference between excited and unexcited regions of an axon
would cause small currents, now called local circuit currents, to flow
between them in such a direction that they stimulate the unexcited region.
Although
excitatory inputs may be seen in the dendrites and/or soma, activation
originates normally only in the soma. Activation in the form of the nerve
impulse (action potential) is first seen in the root of the axon - the initial
segment of the axon, often called the axon hillock. From there it
propagates along the axon. If excitation is initiated artificially somewhere
along the axon, propagation then takes place in both directions from the
stimulus site. The conduction velocity depends on the electric properties and
the geometry of the axon. An important physical property of the membrane is the change in sodium
conductance due to activation. The higher the maximum value achieved by the
sodium conductance, the higher the maximum value of the sodium ion current and
the higher the rate of change in the membrane voltage. The result is a higher
gradient of voltage, increased local currents, faster excitation, and increased
conduction velocity. The decrease in the threshold potential facilitates the
triggering of the activation process. The capacitance of the
membrane per unit length determines the amount of charge required to achieve a
certain potential and therefore affects the time needed to reach the threshold.
Large capacitance values, with other parameters remaining the same, mean a
slower conduction velocity. The velocity also depends
on the resistivity of the medium inside and outside the membrane since these
also affect the depolarization time constant. The smaller the resistance, the
smaller the time constant and the faster the conduction velocity. The
temperature greatly affects the time constant of the sodium conductance; a
decrease in temperature decreases the conduction velocity. The above effects are
reflected in an expression derived by Muler and Markin (1978) using an idealized
nonlinear ionic current function. For the velocity of the propagating nerve
impulse in unmyelinated axon, they obtained
(2.1)
where
v
= velocity of the nerve impulse [m/s]
iNa max
= maximum sodium current per unit length [A/m]
Vth
= threshold voltage [V]
ri
= axial resistance per unit length [W/m]
cm
= membrane capacitance per unit length [F/m]
A myelinated axon (surrounded by the myelin sheath) can produce
a nerve impulse only at the nodes of Ranvier. In these axons the nerve impulse
propagates from one node to another, as illustrated in Figure 2.12. Such a
propagation is called saltatory conduction (saltare, "to dance" in
Latin). The
membrane capacitance per unit length of a myelinated axon is much smaller than
in an unmyelinated axon. Therefore, the myelin sheath increases the conduction
velocity. The resistance of the axoplasm per unit length is inversely
proportional to the cross-sectional area of the axon and thus to the square of
the diameter. The membrane capacitance per unit length is directly proportional
to the diameter. Because the time constant formed from the product controls the
nodal transmembrane potential, it is reasonable to suppose that the velocity
would be inversely proportional to the time constant. On this basis the
conduction velocity of the myelinated axon should be directly proportional to
the diameter of the axon. This is confirmed in Figure 2.13, which shows the
conduction velocity in mammalian myelinated axons as linearly dependent on the
diameter. The conduction velocity in myelinated axon has the approximate value
shown:
v = 6d
(2.2)
where
v
= velocity [m/s]
d
= axon diameter [µm]
Fig. 2.12. Conduction of a nerve impulse in a nerve axon. (A)
continuous conduction in an unmyelinated axon; (B) saltatory conduction in
a myelinated axon.
Fig. 2.13. Experimentally determined conduction velocity of a nerve
impulse in a mammalian myelinated axon as a function of the diameter. (Adapted
from Ruch and Patton, 1982.)
REFERENCES
Berne RM, Levy MN (1993): Physiology, 3rd ed., 1091 pp.
C. V. Mosby, St. Louis.
Bullock TH (1959): Neuron doctrine and electrophysiology.
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Davis LJ, Lorente de Nó R (1947): Contributions to the
mathematical theory of the electrotonus. Stud. Rockefeller Inst. Med.
Res. 131: 442-96.
Elsberg CA (1931): The Edwin Smith surgical papyrus. Ann.
Med. Hist. 3: 271-9.
Ganong WF (1991): Review of Medical Physiology, 15th
ed., Appleton & Lange, Norwalk, Conn.
Guyton AC (1992): Human Physiology and Mechanisms of
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elektrischen Erregung. Pflüger Arch. ges. Physiol. 75: 574-90.)
Hermann L (1905): Lehrbuch der Physiologie, 13th ed.,
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Kandel ER, Schwartz JH (1985): Principles of Neural
Science, Elsevier Publishing, New York.
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