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PHYSICS OF BIOLOGICAL MEMBRANE

Electrolytes are inorganic compounds, where Water is often necessary for

the ionization or molecule split. The dissolved compounds in liquids (water) are
broken down into ions. Ions are atoms, which have accepted more electrons
(negative charges Ion) or lost electrons (positive charges Ion). When solid NaCl is
dissolved in water, water splits the NaCl molecules into free Na + _ and Cl- _ ions,
the solution so formed is called an electrolyte. The ions in electrolyte solution,
generally are more free to move (have a higher mobility) than in solids.

Living tissue is electrically an electrolytic conductor, where both

intracellular and extracellular liquids contain ions free to migrate.

Electrolyte solution has positive and negative ions, if two electrodes with
potential difference are inserted into this electrolyte solution, ion will be attracted
to the electrode with opposite charge. The positively charged ions move toward the
cathode while the negatively charged ions move toward the anode. Therefore an
electric field forms between the two electrodes and such movement of ions
conducts an electric current through the electrolyte solution Figure (1).

Figure (1)

Cell Membrane
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The basic building block of the human body is the living cell, and a
prerequisite for its life is that it is surrounded by an electrolyte solution.
Cells appear as transparent medium when viewed through a light
microscope. In living tissue important communication control is implemented by
hormones and nerves. Some cells are not excitable, for instance the cells of adipose
and connective tissue or blood. They are passive, not under nerve control, and only
weakly polarized. However, nerve, muscle and gland cells are polarized and
excitable; within a 1/1000 second (1 m. second) such cells may react on trigger
signals such as electrical, mechanical or chemical energy.

The excitation of a cell is accompanied by an action potential. The

action potential is the basic bioelectric event and signal source in the body.

The cell is normally surrounded by a membrane, which act as partition to

separate the interior of the cell from its surrounded medium. The membrane is a bi-
layer lipid membrane (BLM) shown in Figure (2). The membrane of a living cell is
a most complex and dynamic system. The structure of the membrane composed of
lipids and proteins with minute pores distributed through the membrane, it is a
major barrier to ion flux, but embedded in the membrane are channels, ion pumps
and...etc.

Figure (2)
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ELECTRICAL PROPERTIES OF CELL MEMBRANE :

MEMBERANE VISCOSITY

The viscosity of a fluid will be constant regardless of the flow field if

the temperature and the density are invariant. Fluids that consist of small
molecules, such as air and water, have low viscosity comparing with blood
viscosity.

Temperature(C°) Water Viscosity (Pa·s)

30 0.797 × 10−3

40 0.653 × 10−3

Temperature(C°) blood Viscosity (Pa·s)

37 3×10−3 to 4×10−3

Blood is a liquid that consists of plasma and particles, such as the red
blood cells. The viscosity of blood thus depends on the viscosity of the plasma, in
combination with the hematocrit.

Physical studies have shown that the membrane exists in a liquid-

crystalline state, where the viscosity of biological membrane ranges from 0.2 poise
and upwards at physiological temperatures of 37C0.

RESISTANCE

The resistor is a circuit component that opposes current flow. Resistance (R)
is measured in units of ohm(Ω).The relation between current (I ) and voltage (V ) is
given by Ohm’s law, which is

V = IR

Where l = length of the conductor.

a = cross-sectional area.
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σ = specific resistance.

Materials that present a very small resistance to current flow are called
conductors. Materials with a very large resistance are called insulators.

In the aqueous environment of the body, salt and various other molecules
dissociate into positive and negative ions. As a result, body fluids are relatively
good conductors of electricity. Still, these fluids are not nearly as conductive as
metals.

The electrical resistivity of the internal fluid is relatively higher than the
external fluid, which is due to its relatively high specific resistance, a smaller
volume and so a narrow cross-sectional area, but the plasma membrane offers the
highest resistance that range between 102 - 105 ohms.cm-2 and consequently limits
the flow of ions through itself. Therefore, it is a far weaker conductor because of
its lipid matrix.

DIELECTRIC

A dielectric is an insulator material, which prevents the movements of

charges from the negative plate to the positive plate through it. Basically, a perfect
dielectric is a substance without free charges. It reduces the intensity of the electric
field between the conductors. The dielectric constant is a main factor, which play
an important role in the design of the capacitor, where the high dielectric constant
enhances the capacitances.
In living cell, membrane capacitance has significant function. Even if the
conductivity of the membrane itself is very low, the membrane is so thin that the
capacitance is very high, that is due to its thickness 7 nm and its dielectric constant
of 3 – 10. This represents a large dielectric strength and it acts as a dielectric, but it
is not a perfect dielectric.

CAPACITANCE

The capacitor is a circuit element that stores electric charges. In its simplest
form it consists of two conducting plates separated by a dielectric (insulator)
shown in Figure (3). In a charged capacitor, positive charges are on one side of the
plate, and negative charges are on the other. Capacitance (C) is measured in farads
(uF.cm-2). The relation between the stored charge (Q), and the voltage across the
capacitor is given by
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C= Q/V

Figure (3)

The intracellular and extracellular fluids are electrolyte solutions contain

electrolyte ions act as conducting plates. A very minute excess of cations
accumulates on the outer surface of the cell membrane and equal number of anions
accumulates on the inner surface. The intracellular and extracellular electrolytes
are well conductive; the cell interior is nearly completely insulated from the
outside by a membrane.

The parallel plate membrane capacitor has a constant and relatively high
capacitance per unit area of the membrane (µF.cm-2) because the membrane is
extremely thin, has relatively high dielectric constant (3 - 10), and the conductive
fluids (outer and inner) offer relatively large surface area towards the membrane.
The parallel-plate membrane capacitor is not perfect capacitors because the
membrane is not perfect dielectric and the ions can diffuse through its pores
leading to dielectric loss and it needs the active membrane transport to maintain its
capacitance.

CELL POLARIZATION (RESTING STATE):

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Cell polarization is generated by the ion pumps, which pump (drive) the
ions against the electrochemical gradient. This energy-consuming mechanism
polarizes the cell so that the interior of excitable cells has a potential about _ 90 mV
with respect to the extracellular electrolytes Figure (4). Such a pump is a molecular
device, embedded in the cell membrane, capable of generating a net electric
current across the membrane. The cell membrane is somewhat permeable to
potassium ions and much less to sodium ions whereas chloride ions can readily
pass through the membrane

Figure (4)

The distribution of ions across the cell membrane and the nature of this
membrane provide the explanation for the membrane potential. The concentration
gradient for K+ facilitates its movement out of the cell via K+ channels, but its
electrical gradient is in the opposite direction (inward). The Na+–K+ pump, pumps
three Na+ out of the cell for every two K+ it pumps in; thus, it also contributes a
small amount to the membrane potential by itself.
Millions of such pumps in one cell membrane polarize the cell to steady
state so that the cell is fully polarized and ready to be triggered. This is the stage
before the action potential and it is known as Resting Membrane Potential.
Where the measuring values of Na+ and K+ concentration are as follow:

Concentration inside Concentration outside

Na+ 15 150 m Mol/ l

K+ 150 m Mol/ l 5.5

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ACTION POTENTIAL

The cell membrane is polarized at rest, with positive charges lined up along
the outside of the membrane and negative charges along the inside. When a surface
of a single axon or section of a cell membrane (nerve or muscle) is excited
effectively and if excitation exceeded firing level either by the flow of ionic current
or by some form of external energy (chemical, thermal, mechanical, electrical…
etc.), changing in membrane characteristics (properties) starts; resistance decreases
and sodium ions enter the cell, thus the charge in the inner surface increases in the
positive direction. Then inward current is formed and this led to decrease the
barrier of the membrane to sodium ions, the sodium channel gate opens and Na +
flows into the cell, the result is rush of sodium ions into the cell and the cell is
depolarized Figure (5).

Figure (5)

The potential crosses the zero line and soon reaches its maximum positive
potential. This potential is known as the action potential and is approximately say
+40 mV. The cell is said to be depolarized and the process of changing from
resting state to the action potential is called depolarization, which is the beginning
of an action potential.

Once the action potential reaches +40 mV, the membrane closes the
+
Na channels, which rapidly enter a closed state in order to block the movement of
sodium ions. In addition, the direction of the electrical gradient for Na + is reversed
during the overshoot because the membrane potential is reversed, and this limits
Na+ influx. The membrane permits some of the potassium ions to leave the cell,
thus the outflow of potassium constitutes an outward current, where the polarity is
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abolished for a brief period and actually reverses (from positive to negative) and
falls rapidly toward the resting level as shown in Figure (6).

Figure (6)

The cell is said to be repolarized and assumes its resting potential when full
repolarization has occurred. The excess sodium ions are actively pumped out of the
cell whereas the excess potassium is actively pumped into the cell.

STANDARD WAVEFORM OF A. P.

In the normal resting state, the potential across the nerve fiber is about
-90mV. Standard waveform is graphical recording of an action potential of a single
nerve fiber, which initiate at the resting potential depolarization, and return to the
resting potential after full repolarized state as shown in Figure (7). Action potential
is a beginning depolarization of the membrane. After the rate of depolarization
increases. The point at which this change in rate occurs is called the firing level or
sometimes the threshold.
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Figure (7) Action potential wave form

Thereafter, the waveform rapidly reaches (zero potential) zero line to

approximately +40 mV. The sharp rise and rapid fall are the spike potential of the
action potential, which is nearly +40 mV.

The ascending phase on the action potential, called the depolarization phase,
is produced by the inward current of sodium ions whereas the descending phase
called the repolarization phase is produced by the outward current of potassium
ions, which represent the reduction of potential from spike potential to resting state
(-90mV).

The total amplitude of the action potential is 130 mV (+40 - (- 90 mV)).That

is defined as the difference between the potential of the depolarized membrane at
the peak of the action potential and the resting membrane potential.
LOCAL RESPONSE

Normally channels have closed gates. A gate can be opened by a voltage

change. The voltage gated is the voltage across the cell membrane that determines
whether channel is opened or remains closed. A polarized cell may suddenly
become depolarized according to the stimulus and threshold intensity, where
threshold intensity varies with the duration; with weak stimuli it is long, and with
strong stimuli it is short.
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The action potential fails to occur if the stimulus is lower than threshold
intensity (around -50mV) and such event is known as local response as shown in
Figure (8).

Figure (8)

Action potential occurs with constant amplitude and form regardless of the
strength of the stimulus if the stimulus is at or above threshold intensity. The local
response can be defined as the event formed in the interval between resting state
and the threshold (firing level).

PERIODS
LATENT PERIOD

The stimulus artifact is followed by an interval (latent period) that ends

with the start of the action potential and corresponds to the time it takes the
impulse to travel along the axon from the site of stimulation to the recording
electrodes as shown in Figure (9).

Figure (9)
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Its duration is proportionate to the distance between the stimulating and

recording electrodes and inversely proportionate to the speed of conduction.
REFRACTORY PERIOD

This refractory period is divided into

absolute refractory period, corresponding to the period from the firing

level until the first-third of the repolarized phase, thus during the depolarization
phase and the first 1/3 of the repolarized phase, the nerve remains absolutely
refractory.

Relative refractory period starts from the end of the first 1/3 of the
repolarized phase, until its last third, during the 2/3 of the repolarized phase nerve
remains relative refractory as shown in Figure (10). During the absolute refractory
period, no stimulus, no matter how strong, will excite the nerve, but during the
relative refractory period, stronger than normal stimuli can cause excitation.

Figure (10)

Transmission of Nerve Action Potential

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Electrical potential exists across the enveloping membrane of living tissue.

Many cells such as nerve, muscles and gland have the ability to transmit a change
in these potentials along their membrane. The transmission can be in either
direction. Muscles fibers can transmit action potentials which result in a
contraction of the muscles.

The transmission of nerve action potentials is brought about by the flow of

ionic current around the nerve fiber. These currents are adequate to depolarize
neighboring cells or adjacent areas of the same cells. As a result a wave of a
depolarization will travel along the membrane without attenuation (damping),
followed by a wave of repolarization. The energy used in propagation does not
from the stimulus, but is released by the nerve fiber along its length. Nerves
normally conduct action potential (impulses) in one direction-toward the central
nervous system in sensory fibers, away from it in motor fibers, but all nerves can
conduct action potential in both directions.

The nerve action potential in one fiber is of constant amplitude and shape
and that characteristics can’t be altered by changing the strength or the quality of
the stimulus. Nerve action potentials have an amplitude of approximately 0.1 V
(100 mV), and a duration of 1 msec. their amplitude is measured between the
inside and the outside of the nerve fibers are of two types: myelinated and non-
myelinted, based on that the transmission can be explained as follow.

(A) Transmission of Nerve A. P. along a non-myelinted nerve fiber

In non-myelinted nerve fibers there is no fatty sheath (myelin) and the

fiber consist of a cylindrical semipermeable membrane which surrounds the axon
of a nerve cell. The membrane, which separates the axoplasm from the external
solution, acts as a barrier and prevents the ions in the external solution from mixing
rapidly with the internal solution. The membrane has a high electrical resistance in
a resting axon and electrical capacity. During the nerve action potential the
conductivity of the membrane increases about 100 times, and sodium and
potassium ions move down their concentration gradients.

In the normal resting state, a potential of about -90 mV exists across the
nerve fiber. When a nerve cell with a long nerve fiber is stimulated at some point,
the stimulus produces local depolarization which will be propagated along the
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nerve fiber. Due to the difference in potential between the adjacent depolarized and
polarized portions of the nerve membrane, some cations flow from the depolarized
to the polarized portion through the axoplasm while some anions flow in opposite
direction. i.e. from the polarized to the depolarized portion through the surrounding
interstitial fluid. These currents are sufficient to stimulate the next inactive portion
of the membrane and consequently, the membrane of the inactive portion gets
depolarized to fire an action potential there as illustrated in Figure (5).

Figure (5) continuous conduction in an unmyelinated nerve

These events are repeated in subsequently parts of the nerve fiber and a
wave of depolarization is propagated along the fiber without attenuation followed
by a wave of repolarization. After a brief period (refractory period), the nerve fiber
becomes capable of transmitting a new action potential.

(B) Transmission of Nerve A. P. along a myelinted nerve fiber

In myelinted nerve fibers the axoplasm of the fiber is shown surrounded by

bands of fatty materials called bands of myelin as shown in Figure (6). The short
gaps between the bands are called the nodes of Ranvier. Myelin is a good insulator,
which enables the action potential to skip from one node to the next and if myelin
is continuous along the nerve, no action potential are possible. Since myelin is
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relatively thick, it has a low capacitance and high resistance against the radial flow
of ions across it.

Figure (6-A&B) saltatory conduction in myelinated nerve

At the nodes, the irritable axon membrane is in direct contact with the
surrounding interstitial fluids.The arrival of an action potential depolarizes the
membrane of a node by enhancing the membrane permeability to sodium ions and
then to potassium ions. Due to the difference in potential between the depolarized
node and the polarized node, some cations flow through the axoplasm from the
depolarized node to the next polarized node and through the interstitial fluid in the
reverse direction. As a result, the membrane of the inactive node gets depolarized
to fire action potential (spike) there, but such formation of action potential can’t
occur in the myelinated bands. The action potential thus, groups from one node to
the next. This process is called salutatory conduction which permits an action
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potential to be transmitted with a higher velocity a long a myelinated nerve than

along a no-myelinated nerve of similar axonal diameter.

Conduction Velocity

In nerve fibers the rate at which an action potential moves down a fiber or is
propagated from cell to cell is called conduction velocity. It varies widely,
depending on nerve diameter, temperature and myelinated. It ranges from 20 to
150 msec. the conduction velocity is higher for myelinated than for non myelinated
and for their nerves. The thinner the nerve fiber, the smaller is the cross-section
area and therefore the higher is the resistance of the axoplasm against the ionic
current. Thus the time needed for the formation of action potential (spike) in active
region is prolonged therefore, the speed of the spike falls and the duration
increases in proportion to the fall in the fiber diameter. Larger nerves conduct
action potential faster than then nerves. In muscles, the conduction is much slower
and contraction follows the development of action potential. The rate of travel of
action potential through the muscle is slower with 0.2 to o.4 msec. on the average.
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