Biomedical Instrumentation

UNIT - I: Bio-Potential Signals and Electrodes

Basic Medical Instrumentation System

The primary purpose of medical instrumentation is to measure or determine the

presence of some physical quantity that may some way assist the medical personnel to
make better diagnosis and treatment. Accordingly, many types of instrumentation systems
are presently used in hospitals and other medical facilities. The majority of the instruments
are electrical or electronic systems, although mechanical systems such as ventilators or
spirometers are also employed. Because of the predominantly large number of electronic
systems used in medical practice, the concepts explained hereafter are mostly related to
electronic medical instruments.
Certain characteristic features, which are common to most instrumentation systems,
are also applicable to medical instrumentation systems.

Measurand: The physical quantity or condition that the instrumentation system measures
are called the measurand. The source for the measurand is the human body which generates
a variety of signals. The measurand may be on the surface of the body (electrocardiogram
potential) or it may be blood pressure in the chambers of the heart.

Transducer/Sensor: A transducer is a device that converts one form of energy to another.

Because of the familiar advantages of electric and electronic methods of measurement, it is
the usual practice to convert into electrical quantities all non-electrical phenomenon
associated with the measurand with the help of a transducer.
Signal Conditioner: Converts the output of the transducer into an electrical quantity
suitable for operation of the display or recording system. Signal conditioners may vary in
complexity from a simple resistance network or impedance matching device to multi-stage
amplifiers and other complex electronic circuitry.

Display System: Provides a visible representation of the quantity as a displacement on a

scale, or on the chart of a recorder, or on the screen of a cathode ray tube or in numerical
form. Although, most of the displays are in the visual form, other forms of displays such as
audible signals from alarm or foetal Doppler ultrasonic signals are also used.

Alarm System with upper and lower adjustable thresholds to indicate when the measurand
goes beyond preset limits.
Data Storage to maintain the data for future reference. It may be a hard copy on a paper or
on magnetic or semiconductor memories.
Data Transmission using standard interface connections so that information obtained may
be carried to other parts of an integrated system or to transmit it from one location to
another.

Sources of Biomedical Signals

Biomedical signals are those signals (phenomenon that conveys information) which are
used primarily for extracting information on a biological system under investigation. The
process of extracting information could be as simple as feeling the pulse of a person on the
wrist or as complex as analyzing the structure of internal soft tissues by an ultrasound
scanner. Biomedical signals originate from a variety of sources.

Bio signals: Biological signals or biosignals are space, time or space-time records of a
biological event such as a beating heart or a contracting muscle. The electrical, chemical
and mechanical activity that occurs during this biological event often produces signals that
can be measured and analyzed. The Biological signals are very much useful in medical
diagnostics. These signals may be classified into Bioelectric Signals and Non-Bioelectric
Signals.
Bioelectric Signals: These are unique to the biomedical systems. They are generated by
nerve cells and muscle cells. Their basic source is the cell membrane potential which under
certain conditions may be excited to generate an action potential. The electric field
generated by the action of many cells constitutes the bio-electric signal. The most common
examples of bioelectric signals are the ECG (electrocardiographic), EEG
(electroencephalographic), EMG (Electromyogram), EOG (Electrooculogram), and ERG
(Electrorentiogram) signals.
Bioacoustic Signals: The measurement of acoustic signals created by many biomedical
phenomena provides information about the underlying phenomena. The examples of such
signals are: flow of blood in the heart, through the hearts valves and flow of air through the
upper and lower airways and in the lungs which generate typical acoustic signal.
Biomechanical Signals: These signals originate from some mechanical function of the
biological system. They include all types of motion and displacement signals, pressure and
flow signals etc.
Biochemical Signals: The signals which are obtained as a result of chemical measurements
from the living tissue or from samples analyzed in the laboratory. The examples are
measurement of partial pressure of carbon-dioxide (pCO2), partial pressure of oxygen
(pO2) and concentration of various ions in the blood.
Biomagnetic Signals: Extremely weak magnetic fields are produced by various organs
such as the brain, heart and lungs. The measurement of these signals provides information
which is not available in other types of bio-signals such bio-electric signals. A typical
example is that of magneto-encephalograph signal from the brain.
Bio-optical Signals: These signals are generated as result of optical functions of the
biological systems, occurring either naturally or induced by the measurement process. For
example, blood oxygenation may be estimated by measuring the transmitted/back scattered
light from a tissue at different wavelengths.
Bio-impedance Signals: The impedance of the tissue is a source of important information
concerning its composition, blood distribution and blood volume etc. The measurement of
galvanic skin resistance is a typical example of this type of signal. The bio-impedance
signal is also obtained by injecting sinusoidal current in the tissue and measuring the
voltage drop generated by the tissue impedance. The measurement of respiration rate based
on bio-impedance technique is an example of this type of signals.

What is Biosignals?

A biosignals is any signal in living beings that can be continually measured and monitored.
The term biosignals is often used to refer to bioelectrical signals, but it may refer to both
electrical and non-electrical signals. The usual understanding is to refer only to time-
varying signals, although spatial parameter variations are sometimes subsumed as well.

Biological signals, or biosignals, are space, time, or space-time records of a biological

event such as a beating heart or a contracting muscle. The electrical, chemical, and
mechanical activity that occurs during this biological event often produces signals that can
be measured and analyzed.

A biosignals can be defined as a physiological phenomenon, a body variable that can be

measured and monitored. Since the number of physiological mechanisms is nearly
unlimited, the diversity of biosignals is huge. This can also be justified by the fact that there
are many ways to classify the biosignals:
 PERMANENT/INDUCED (INTRINSIC/EXTRINSIC TO BODY)
 Static/Dynamic
 Origin
As referred the variety of biosignals is nearly unlimited, this makes a unique classification
of biosignals impossible. Their classification is based on their characteristics.
Classification
Intrinsic/Extrinsic to body
This first method takes the existence of biosignals as a way to classify them, dividing the
biosignals into:
 Permanent Biosignals
This kind of Biosignals exist without any excitation from outside body and are always
present in the Human Body because source is inside the body. One example is the
electrocardiographic signal (ECG) induced by electrical heart muscle excitation with the
peaks P-Q-R-T-S.
 Induced Biosignals
This group of biosignals includes biosignals that are artificially induced. In contrast with
the permanent biosignals this one’s exist only during the excitation. It means that, when the
artificial induction is over the induced biosignal decays with a time constant determined by
the body properties. One example is electric plethsysmography, here an artificial current is
induced in the tissue.
Dynamic
The second method takes in consideration the dynamic nature of the biosignal according to:
 Static Biosignal
Static biosignals carry information during their steady-state lever which may show slow
changes over the time. For example the body temperature, which shows slightly, changes
during the day, that's why we can consider it a static biosignal.
 Dynamic Biosignal
Dynamic biosignals show big changes during time, for example the heart rate. The course
of the heart rate represents a highly dynamic biosignal.
Origin
The last method is using the origin of the biosignal as a basis for their classification, here
are some examples:
 Electric Biosignals
 Magnetic Biosignals
 Mechanic Biosignals
 Optic Biosignals
 Acoustic Biosignals
 Chemical Biosignals
 Thermal Biosignal
Active Vs Passive

Biosignals can also be divided in two main groups according to their source for
measurement: Active and Passive.
Active
These are biosignals where the energy source for measurement is the patient himself. Here
we have two types of "sub" biosignals:
 Electrical Biosignals: ECG; EEG; EMG.
 Non-electrical biosignals: Thermography and pH for example.
Passive
These ones, the energy source for measurement is not the patient e.g. wrist oximeter.
== Electric Biosignals == WHAT HAS THIS GOT TO DO WITH CHARACTERISTICS
OF SIGNALS???
Knowing the different signals from the brain and from other parts of the body is very
important to understand the reason of many physiological and pathological functions of
these same ones. Electric biosignals can be defined as a change in the electric current across
a specialized tissue, organ or cell like the nervous system for example. Some examples of
electric biosignals are:
 Electrocardiogram (ECG)
 Electroencephalogram (EEG)
 Electromyogram (EMG)
ECG
The electrocardiogram is a graphic which is produced by an electrocardigraph. This device
records the heart activity over the time. When the electrical waves which cause the heart
muscles to contract pass-through the body, they can be measured by the electrodes placed
on the patient skin, providing a view of the heart muscle activity. A typical ECG tracing is
a cycle of three entities.

Biosignal Characteristics

The term biosignal refers to all the signals that are being generated in all living organisms.
They can also be described as the spaces, time or the space - time records of a biological
event inside the body such as the heart beating or the contraction of the muscles so all the
electrical, chemical and mechanical activities that happens during these events produce
signals which can be measured and analyzed .these signals are so useful to understand the
biological events of the body which can help in medical diagnosis.
Physiological origins of Biosignals
Bioelectric signals
Are generated by nerves and muscles tissues as the result of the changes in the electric
currents which are produced by the sum potential differences across the tissues and organs.
Best known example is the Electrocardiography.
Biomagnatic signals
Different organs such as heart, lungs, brain generate weak magnetic field. It measured from
activities which are linked to electric field from an organ.
Example: magnetcardiography.
Biochemical signals
Signals contain information about the changes in concentration of various chemical agents
in the body.
For example it determines the level of glucose.
Biomechanical signals
Produced by the mechanical functions of biological signals such as motion and
displacement, pressure.
Example: blood pressure measurements.
Bioacoustic signals
They are special subset of biomechanical signals that involve in vibration basically motion.
Example: respiratory system and muscles generates this kind of signals.
Biooptical signals
They are generated by the optical or light induced attributes of biological systems. They
may occur naturally or induced.

Biosignal Characteristics
Biosignals can be classified according to various characteristics:
Waveform shape
It is the shape of a signal. The waveform represents the variation of a voltage or current of a
signal over time in graph.
Statisticals structures of the signals
Temporal properties
They are the changes in the signals according to time.
Classification of Biosignals

Continuous Signals

Defined over a continuum of time or space and are described by continuous variable
functions. They are produced by biological phenomena:
 Voltages measurements from the heart
 Arterial blood measurements
 Measurements of electrical activities from the brain

Discrete signals
Discrete signals define only at subset of regularly spaced points in time and-or space.
Continuous signals from human body are converted to discrete signals by a process called
sampling and they can be analyzed and interpreted by a computer.

Organization of cell
Cells are the basic building blocks of living things. The human body is composed of
trillions of cells, all with their own specialised function. 

Illustration showing the structures of an animal cell

 Cells are the basic structures of all living organisms.
 Cells provide structure for the body, take in nutrients from food and carry out
important functions.
 Cells group together to form tissues, which in turn group together to form organs,
such as the heart and brain.
 Our cells contain a number of functional structures called organelles
 These organelles carry out tasks such as making proteins, processing chemicals and
generating energy for the cell.
 The nucleus is based at the centre of the cell and is the ‘control room’ for the cell.
 The genome is found within the nucleus.

Organization of cells in the human body

Cells are organized into tissues, and tissues form organs. Organs are organized into organ
systems such as the skeletal and muscular systems.

A cell is the smallest independently functioning unit of a living organism. Even bacteria,
which are extremely small, independently-living organisms, have a cellular structure. Each
bacterium is a single cell. All living structures of human anatomy contain cells, and almost
all functions of human physiology are performed in cells or are initiated by cells.
A human cell typically consists of flexible membranes that enclose cytoplasm, a water-
based cellular fluid together with a variety of tiny functioning units called organelles. In
humans, as in all organisms, cells perform all functions of life. A tissue is a group of many
similar cells (though sometimes composed of a few related types) that work together to
perform a specific function. An organ is an anatomically distinct structure of the body
composed of two or more tissue types. Each organ performs one or more specific
physiological functions. An organ system is a group of organs that work together to
perform major functions or meet physiological needs of the body.

Nernst equation of membrane

In electrochemistry, the Nernst equation is an equation that relates the reduction potential of

a reaction (half-cell or full cell reaction) to the standard electrode potential, temperature,
and activities (often approximated by concentrations) of the chemical species undergoing
reduction and oxidation. It was named after Walther Nernst, a German physical
chemist who formulated the equation.

Nernst potential

The Nernst equation has a physiological application when used to calculate the potential of
an ion of charge z across a membrane. This potential is determined using the concentration
of the ion both inside and outside the cell:

E= {RT}/{zF}} * ln{ion outside cell}/{ion inside cell}

=2.3026 * {RT}/{zF}* log10{ion outside cell}/{ion inside cell}

When the membrane is in thermodynamic equilibrium (i.e., no net flux of ions), and if the
cell is permeable to only one ion, then the membrane potential must be equal to the Nernst
potential for that ion.

Where

Em is the membrane potential (in volts, equivalent to joules per coulomb),

Pion is the permeability for that ion (in meters per second),
[ion]out is the extracellular concentration of that ion (in moles per cubic meter, to
match the other SI units, though the units strictly don't matter, as the ion
concentration terms become a dimensionless ratio),
[ion]in is the intracellular concentration of that ion (in moles per cubic meter),
 R is the ideal gas constant (joules per Kelvin per mole),
T is the temperature in Kelvin’s,
F is Faraday's constant (coulombs per mole).
z is the valence of the ionic species. For example, z is +1 for Na+, +1 for K+, +2 for
Ca2+, −1 for Cl−, etc.

Resting and Action potentials

In physiology, an action potential (AP) occurs when the membrane potential of a

specific cell location rapidly rises and falls  this depolarization then causes adjacent
locations to similarly depolarize. Action potentials occur in several types of animal cells,
called excitable cells, which include neurons, muscle cells, endocrine cells and in
some plant cells.
In neurons, action potentials play a central role in cell-to-cell communication by providing
for or with regard to saltatory conduction, assisting the propagation of signals along the
neuron's axon toward synaptic boutons situated at the ends of an axon; these signals can
then connect with other neurons at synapses, or to motor cells or glands. In other types of
cells, their main function is to activate intracellular processes. In muscle cells, for example,
an action potential is the first step in the chain of events leading to contraction. In beta
cells of the pancreas, they provoke release of insulin. Action potentials in neurons are also
known as "nerve impulses" or "spikes", and the temporal sequence of action potentials
generated by a neuron is called its "spike train". A neuron that emits an action potential, or
nerve impulse, is often said to "fire".
Action potentials are generated by special types of voltage-gated ion channels embedded in
a cell's plasma membrane. These channels are shut when the membrane potential is near the
(negative) resting potential of the cell, but they rapidly begin to open if the membrane
potential increases to a precisely defined threshold voltage, depolarising the transmembrane
potential. When the channels open, they allow an inward flow of sodium ions, which
changes the electrochemical gradient, which in turn produces a further rise in the membrane
potential towards zero. This then causes more channels to open, producing a greater electric
current across the cell membrane and so on. The process proceeds explosively until all of
the available ion channels are open, resulting in a large upswing in the membrane potential.
The rapid influx of sodium ions causes the polarity of the plasma membrane to reverse, and
the ion channels then rapidly inactivate. As the sodium channels close, sodium ions can no
longer enter the neuron, and they are then actively transported back out of the plasma
membrane. Potassium channels are then activated, and there is an outward current of
potassium ions, returning the electrochemical gradient to the resting state. After an action
potential has occurred, there is a transient negative shift, called the after hyper polarization.
In animal cells, there are two primary types of action potentials. One type is generated
by voltage-gated sodium channels, the other by voltage-gated calcium channels. Sodium-
based action potentials usually last for under one millisecond, but calcium-based action
potentials may last for 100 milliseconds or longer. In some types of neurons, slow calcium
spikes provide the driving force for a long burst of rapidly emitted sodium spikes.
In cardiac muscle cells, on the other hand, an initial fast sodium spike provides a "primer"
to provoke the rapid onset of a calcium spike, which then produces muscle contraction.

Overview

Nearly all cell membranes in animals, plants and fungi maintain a voltage difference

between the exterior and interior of the cell, called the membrane potential. A typical
voltage across an animal cell membrane is −70 mV. This means that the interior of the cell
has a negative voltage relative to the exterior. In most types of cells, the membrane
potential usually stays fairly constant. Some types of cells, however, are electrically active
in the sense that their voltages fluctuate over time. In some types of electrically active cells,
including neurons and muscle cells, the voltage fluctuations frequently take the form of a
rapid upward spike followed by a rapid fall. These up-and-down cycles are known as action
potentials. In some types of neurons, the entire up-and-down cycle takes place in a few
thousandths of a second. In muscle cells, a typical action potential lasts about a fifth of a
second. In some other types of cells and plants, an action potential may last three seconds
or more.
The electrical properties of a cell are determined by the structure of the membrane that
surrounds it. A cell membrane consists of a lipid bi-layer of molecules in which larger
protein molecules are embedded. The lipid bi-layer is highly resistant to movement of
electrically charged ions, so it functions as an insulator. The large membrane-embedded
proteins, in contrast, provide channels through which ions can pass across the membrane.
Action potentials are driven by channel proteins whose configuration switches between
closed and open states as a function of the voltage difference between the interior and
exterior of the cell. These voltage-sensitive proteins are known as voltage-gated ion
channels.
Process in Typical Neuron

All cells in animal body tissues are electrically polarized – in other words, they maintain a
voltage difference across the cell's plasma membrane, known as the membrane potential.
This electrical polarization results from a complex interplay between protein structures
embedded in the membrane called ion pumps and ion channels. In neurons, the types of ion
channels in the membrane usually vary across different parts of the cell, giving
the dendrites, axon, and cell body different electrical properties. As a result, some parts of
the membrane of a neuron may be excitable (capable of generating action potentials),
whereas others are not. Recent studies have shown that the most excitable part of a neuron
is the part after the axon hillock (the point where the axon leaves the cell body), which is
called the initial segment, but the axon and cell body are also excitable in most cases.

Each excitable patch of membrane has two important levels of membrane potential:
the resting potential, which is the value the membrane potential maintains as long as
nothing perturbs the cell, and a higher value called the threshold potential. At the axon
hillock of a typical neuron, the resting potential is around –70 millivolts (mV) and the
threshold potential is around –55 mV. Synaptic inputs to a neuron cause the membrane
to depolarize or hyperpolarize; that is, they cause the membrane potential to rise or fall.
Action potentials are triggered when enough depolarization accumulates to bring the
membrane potential up to threshold. When an action potential is triggered, the membrane
potential abruptly shoots upward and then equally abruptly shoots back downward, often
ending below the resting level, where it remains for some period of time. The shape of the
action potential is stereotyped; this means that the rise and fall usually have approximately
the same amplitude and time course for all action potentials in a given cell. (Exceptions are
discussed later in the article). In most neurons, the entire process takes place in about a
thousandth of a second. Many types of neurons emit action potentials constantly at rates of
up to 10–100 per second. However, some types are much quieter, and may go for minutes
or longer without emitting any action potentials.

Resting Potential

The relatively static membrane potential of quiescent cells is called the resting membrane

potential (or resting voltage), as opposed to the specific dynamic electrochemical
phenomena called action potential and graded membrane potential.
Apart from the latter two, which occur in excitable cells (neurons, muscles, and some
secretory cells in glands), membrane voltage in the majority of non-excitable cells can also
undergo changes in response to environmental or intracellular stimuli. The resting potential
exists due to the differences in membrane permeabilities for potassium, sodium, calcium,
and chloride ions, which in turn result from functional activity of various ion channels, ion
transporters, and exchangers. Conventionally, resting membrane potential can be defined as
a relatively stable, ground value of transmembrane voltage in animal and plant cells.
The typical resting membrane potential of a cell arises from the separation
of potassium ions from intracellular, relatively immobile anions across the membrane of the
cell. Because the membrane permeability for potassium is much higher than that for other
ions, and because of the strong chemical gradient for potassium, potassium ions flow from
the cytosol into the extracellular space carrying out positive charge, until their movement is
balanced by build-up of negative charge on the inner surface of the membrane. Again,
because of the high relative permeability for potassium, the resulting membrane potential is
almost always close to the potassium reversal potential. But in order for this process to
occur, a concentration gradient of potassium ions must first be set up. This work is done by
the ion pumps/transporters and/or exchangers and generally is powered by ATP.
In the case of the resting membrane potential across an animal cell's plasma membrane,
potassium (and sodium) gradients are established by the Na+/K+-ATPase (sodium-
potassium pump) which transports 2 potassium ions inside and 3 sodium ions outside at the
cost of 1 ATP molecule. In other cases, for example, a membrane potential may be
established by acidification of the inside of a membranous compartment (such as the proton
pump that generates membrane potential across synaptic vesicle membranes).
Generation of Resting Potential

Cell membranes are typically permeable to only a subset of ions. These usually include
potassium ions, chloride ions, bicarbonate ions, and others. To simplify the description of
the ionic basis of the resting membrane potential, it is most useful to consider only one
ionic species at first, and consider the others later. Since trans-plasma-membrane potentials
are almost always determined primarily by potassium permeability, that is where to start.
 Panel 1 of the diagram shows a diagrammatic representation of a simple cell where a
concentration gradient has already been established. This panel is drawn as if the
membrane has no permeability to any ion. There is no membrane potential because
despite there being a concentration gradient for potassium, there is no net charge
imbalance across the membrane. If the membrane were to become permeable to a type
of ion that is more concentrated on one side of the membrane, then that ion would
contribute to membrane voltage because the permeant ions would move across the
membrane with net movement of that ion type down the concentration gradient. There
would be net movement from the side of the membrane with a higher concentration of
the ion to the side with lower concentration. Such a movement of one ion across the
membrane would result in a net imbalance of charge across the membrane and a
membrane potential. This is a common mechanism by which many cells establish a
membrane potential.
 In panel 2 of the diagram, the cell membrane has been made permeable to potassium
ions, but not the anions (An−) inside the cell. These anions are mostly contributed by
protein. There is energy stored in the potassium ion concentration gradient that can be
converted into an electrical gradient when potassium (K +) ions move out of the cell.
Note that potassium ions can move across the membrane in both directions but by the
purely statistical process that arises from the higher concentration of potassium ions
inside the cell, there will be more potassium ions moving out of the cell. Because there
is a higher concentration of potassium ions inside the cells, their random molecular
motion is more likely to encounter the permeability pore (ion channel) that is the case
for the potassium ions that are outside and at a lower concentration. An internal K + is
simply "more likely" to leave the cell than an extracellular K+ is to enter it. It is a matter
of diffusion doing work by dissipating the concentration gradient. As potassium leaves
the cell, it is leaving behind the anions. Therefore, a charge separation is developing as
K+ leaves the cell. This charge separation creates a transmembrane voltage. This
transmembrane voltage is the membrane potential. As potassium continues to leave the
cell, separating more charges, the membrane potential will continue to grow. The
 length of the arrows (green indicating concentration gradient, red indicating voltage),
represents the magnitude of potassium ion movement due to each form of energy. The
direction of the arrow indicates the direction in which that particular force is applied.
Thus, the building membrane voltage is an increasing force that acts counter to the
tendency for net movement of potassium ions down the potassium concentration
gradient.
 In Panel 3, the membrane voltage has grown to the extent that its "strength" now
matches the concentration gradients. Since these forces (which are applied to K +) are
now the same strength and oriented in opposite directions, the system is now
in equilibrium. Put another way, the tendency of potassium to leave the cell by running
down its concentration gradient is now matched by the tendency of the membrane
voltage to pull potassium ions back into the cell. K + continues to move across the
membrane, but the rate at which it enters and leaves the cell are the same, thus, there is
no net potassium current. Because the K+ is at equilibrium, membrane potential is
stable, or "resting" (EK).
The resting voltage is the result of several ion-translocating enzymes
(uniporters, cotransporters, and pumps) in the plasma membrane, steadily operating in
parallel, whereby each ion-translocator has its characteristic electromotive force (= reversal
potential = 'equilibrium voltage'), depending on the particular substrate concentrations
inside and outside (internal ATP included in case of some pumps).
H+ exporting ATPase render the membrane voltage in plants and fungi much more negative
than in the more extensively investigated animal cells, where the resting voltage is mainly
determined by selective ion channels.
In most neurons the resting potential has a value of approximately −70 mV. The resting
potential is mostly determined by the concentrations of the ions in the fluids on both sides
of the cell membrane and the ion transport proteins that are in the cell membrane. How the
concentrations of ions and the membrane transport proteins influence the value of the
resting potential is outlined below.
The resting potential of a cell can be most thoroughly understood by thinking of it in terms
of equilibrium potentials. In the example diagram here, the model cell was given only one
permeant ion (potassium). In this case, the resting potential of this cell would be the same
as the equilibrium potential for potassium.
However, a real cell is more complicated, having permeabilities to many ions, each of
which contributes to the resting potential. To understand better, consider a cell with only
two permeant ions, potassium, and sodium. Consider a case where these two ions have
equal concentration gradients directed in opposite directions, and that the membrane
permeabilities to both ions are equal. K+ leaving the cell will tend to drag the membrane
potential toward EK. Na+ entering the cell will tend to drag the membrane potential toward
the reversal potential for sodium ENa. Since the permeabilities to both ions were set to be
equal, the membrane potential will, at the end of the Na +/K+ tug-of-war, end up halfway
between ENa and EK. As ENa and EK were equal but of opposite signs, halfway in between is
zero, meaning that the membrane will rest at 0 mV.
Note that even though the membrane potential at 0 mV is stable, it is not an equilibrium
condition because neither of the contributing ions is in equilibrium. Ions diffuse down their
electrochemical gradients through ion channels, but the membrane potential is upheld by
continual K+ influx and Na+ efflux via ion transporters. Such situation with similar
permeabilities for counter-acting ions, like potassium and sodium in animal cells, can be
extremely costly for the cell if these permeabilities are relatively large, as it takes a lot
of ATP energy to pump the ions back. Because no real cell can afford such equal and large
ionic permeabilities at rest, resting potential of animal cells is determined by predominant
high permeability to potassium and adjusted to the required value by modulating sodium
and chloride permeabilities and gradients.
In a healthy animal cell Na+ permeability is about 5% of the K+ permeability or even less,
whereas the respective reversal potentials are +60 mV for sodium (ENa)and −80 mV for
potassium (EK). Thus the membrane potential will not be right at EK, but rather depolarized
from EK by an amount of approximately 5% of the 140 mV difference between EK and ENa.
Thus, the cell's resting potential will be about −73 mV.
In a more formal notation, the membrane potential is the weighted average of each
contributing ion's equilibrium potential. The size of each weight is the relative conductance
of each ion. In the normal case, where three ions contribute to the membrane potential:
Em = (gk+ / gtot )* Ek+ + (gNa+ / gtot )* ENa+ + (gcl- / gtot )* Ecl-
where
 Em is the membrane potential, measured in volts
 EX is the equilibrium potential for ion X, also in volts
 gX/gtot is the relative conductance of ion X, which is dimensionless
 gtot is the total conductance of all permeant ions in arbitrary units (e.g. siemens for
electrical conductance), in this case gK+ + gNa+ + gCl−