THE ELECTROCARDIOGRAM

For

60 years physicians have used this record of the electrical

activity of the heart for diagnosis. Ne,v investigations show ho"r

it

is related to the electrical events that co-ordinate the heart

by

or more than half a century physi

cians have been recording varia
tions in electrical potential on the
surface of the body and using them to
make increasingly complete and reliable
diagnoses of conditions of the heart. As
often happens in medicine, practice has
moved ahead faster than research.
Physicians can read electrocardiograms
with confidence because thousands of
these records have been correlated with
other findings in the clinic and in autop
sy. In the past this fund of empirical
knowledge has concealed a meager un
derstanding of electrical activity inside
the heart.
Now, it appears, understanding has
begun to overtake practice. Several
groups of investigators using, among
other ingenious techniques, electrodes
fine enough to measure potentials inside
a single cell have arrived at an electro
chemical explanation for the function of
both nerve and muscle [see "How Cells
Communicate," by Bernhard Katz;
September]. In our
laboratory at the University of Wash
ington School of Medicine we have de
veloped a fine multitip extracellular elec
trode with which we have been observ
ing electrical events within the muscle
of the heart. Our studies are b'acing the
pathway of the wave of electrical excita
tion that controls and co-ordinates the
rhythmic contractions of heart muscle.
What contribution such direct study of
cardiac potentials can make to diagnostic
electrocardiography is not yet clear. But
it considerably enhances the picture of
the physiological events that give rise to
the electrocardiogram.
During the recording of a conven
tional electrocardiogram the body acts as
a volume conductor, as though it were a
container of salt solution. Such a conduc
tor extends the potential field around a

ENTIFIC AMERICAN,

Allen M. Scher

current source in a manner analogous to

the extension of a magnetic field around
a magnet. Just as a piece of iron that is
not in direct contact with the magnet is
attracted or repelled by the proximity
of a magnetic pole, so distant recording
electrodes, affixed to the skin with a
conducting paste, respond to the poten
tial field set up by the action of the heart.
A typical electrocardiogram shows a
characteristic potential change in time.
The various waves are known respective
ly as the P wave, the QRS complex and
the T wave, after the designations given
them near the turn of the century by
Willem Einthoven, the Dutch physician
who devised a galvanometer sensitive
enough to permit routine recording of
the electrocardiogram.

----+-3rT-

AORTA

SCI

The onset of the wave closely fol

lows the firing of the pacemaker, a knot
of highly specialized muscle cells that
sets the rhythm of the heart. The P wave
is produced by the movement of a wave
of electrical activity through the auricles,
or atria: the two thin-walled chambers
at the top of the heart. A single mass of
muscle forms the outer walls of these
chambers, and the inner space is palti
tioned by a sheet of muscle called the in
teratrial septum. Electrical activity al
ways precedes mechanical contraction,
and almost immediately after the wave
of electrical activity moves through a
portion of atrial muscle, the muscle con
tracts. During contraction the blood con
tained in the atria Rows into the ventri
cles: the lower chambers of the heart.

n'------

PULMONARY VEINS

PULMONARY ARTERY
INTERATRIAL SEPTUM
RIGHT ATRIUM
CON DUC TI ON
SYS TE M

------f-------+-+
-------+---"

1-----

LEFT ATRIUM

;-"-1-",-"-'-'-,1------+"'----

ATRIOVENTRICULAR

VENA

VALVES

LEFT VENTRICLE

RIGHT

INTERIOR OF HUMAN HEART

is shown in this cuta.way diagram. Anatomical features

mentioned in the article are labeled. Conduction system consists of specialized muscle.

132
1961 SCIENTIFIC AMERICAN, INC

From the right atrium oxygen-poor

venous blood flows into the right ven
tricle; simultaneously from the left
atrium oxygen-rich blood from the lungs
moves into the left ventricle. The atria
are separated from the ventricles by a
ring of electrically inert connective tissue
that contains the atrioventricular valves.
After the P wave there intervenes a
pause of
milliseconds. During this
interval the atrioventricular node and the
attached conducting bundles, which fur
nish the only electrical connection be
tween the upper and lower chambers of
the heart, carry the wave of excitation to
the ventricles. The electrocardiogram
now records the QRS complex. This is
produced by the electrical activity of the
massive muscles of the ventricular walls
and of the interventricular septum. This
electrical activity leads to contraction of
the two ventricles; the right ventricle dis
charges venous blood into the pulmonary
artery and the left ventricle discharges
arterial blood into the aorta for distribu
tion to the systemic arteries. Then fol
lows the T wave, which marks the re
covery of the ventricies for the next
cycle.

80

he potential changes recorded by

electrocardiogram represent the
summation of the electrical activity of
thousands of individual cardiac-muscle
cells. As in all muscle cells, the contrac
tion of these cells is initiated by potential
changes that occur across the cell mem
brane. Between the interior of the rest-

T the

ing cell and its external environment

there is a difference in electrical po
tential that is set up by a difference
in the concentration of ions in the
interior of the cell and in the fluids
that bathe the cells. Inside the cel! the
concentration of sodium is low and that
of potassium high with respect to the
concentrations of these ions outside. A
chemical "pump" somehow maintains
the concentration gradients and a conse
quent potential of
'millivolts across
the membrane of the resting cell; the in
side of the cell is negative with respect
to the outside [see "How Things Get
into Cells," by Heinz Holter;
September]. When a cell
goes into the active, or depolarized, state,
there is a rush of ions across the mem
brane and the inside now becomes about
millivolts positive with respect to the
outside. How this "depolarization" of the
membrane triggers mechanical contrac
tion remains unexplained.
When any cell in the heart depo
larizes, current flows into it from ad
jacent cells. The current flow lowers
the resting potential of adjacent cells,
and these cells thereupon depolarize
explosively. Thus, like a flame sweeping
through gunpowder, a wave of excita
tion sweeps through the entire mass of
muscle. Excitable cells, unlike grains of
gunpowder, spontaneously return to the
polarized resting condition and are ready
to become active again. Nerve cells
spend less than
second in the de
polarized state after firing. Heart cells

80

SCIENTIFIC

AMERICAN,
30

.001

are comparatively slow to recover; they

remain in the depolarized state for a
quarter of a second to half a second be
fore returning to the polarized, resting
and relaxed state.
From the point of view of the record
ing elech'ode of an electrocardiograph
each segment of heart muscle acts as if
it were a large single cell. A resting cell
-and resting tissue-induces no poten
tial at the electrode. Generally speaking,
the influence that an aggregation of
charges exerts on an electrode is a func
tion of the distance to the electrode. The
positive charges on the outer surface
of the membrane' closer to the electrode
obviously exert a larger influence than
do the negative charges on the inner sur
face of the same portion of membrane.
But the negative charges on the inner
surface of the membrane on the opposite
side of the cell are closer to the electrode
than the positive charges on the outer
surface of this portion of membrane.
The positive and negative charges in
resting tissue therefore cancel each
other.
The movement of a wave of activity
through heart tissue causes potential
changes at the electrodes. As the wave
of activity starts, an electrode near its
origin registers a negative potential rep
resenting the combined influence of the
negative charges accumulating outside
the depolarized muscle cells closest to it
and the negative charges inside the cells
in the still-resting tissue beyond. Con
versely, an electrode near the termina-

'\

2
T

.-

Q V- \

.4

.8

1.2

1.6

-u
2

TIME (SECONDS)

NORMAL ELECTROCARDIOGRAM shows well-defined P wave,

which brings on contraction of atria, followed by QRS complex,

2.4

/L

2.8

3.2

3.i6

which causes contraction of ventr.icles. Colored lines on time scale

show duration of atrial and ventricular contractions respectively.

133
1961 SCIENTIFIC AMERICAN, INC

tion of activity registers a positive poten

tial set up by positive charges outside the
cells of the resting tissue closer to it
and the positive charges inside the more
distant cells that have been fired by the
approaching wave. An electrode affixed
to the skin or planted directly in the
heart thus "sees" the movement of a
wave of activity as a sequence of poten
tial changes. It records receding activity
as a negative potential and approaching
activity as a positive potential [see illus
trations on opposite page]. From the
shape of the potential recorded at any
given point on the body, therefore, one
may determine the direction in which
the wave of activity traveled through the
heart with respect to that point.
The electrocardiograph, like any other
electrical amplifier, has two input elec
trodes. Its output is a record of the
difference in potential between them.
For many records it is necessary to null
one of the two inputs, to make it effec
tively indifferent to potential change.
This is accomplished by a network that
connects electrodes placed at three ex
tremities of the body to a single side of
the input amplifier. The potentials at

I-------"
I
I
I
I
I
I
I
I

>

I
I

he shapes of the waves recorded on

surface of the body from the
thin-walled atria were sufficiently clear
to enable the English physiologist Sir
Thomas Lewis to plot the pathway of
atrial activity in
Later work in
other laboratories has not significantly
modified his conclusions. The spontane
ous firing of the pacemaker cells located
on the right atrium induces the depolar
ization of the nearest resting cells, and
the flow of current into these fires the
next nearest cells in turn. The boundary
between resting and depolarized cells
thus sweeps through the entire atrium in
about
milliseconds, at a speed of ap
proximately
meter per second. Mov
ing roughly parallel to the free walls of
the atria and the axis of the interatrial
septum, the wave travels in a plane away
from the right ann and toward the left
leg with little motion perpendicular to
this plane [see top illustration on page
136]. The smooth and orderly motion,

T- the

1914.

80

.4

TIME (SECONDS)

these three sites are thereby averaged to

approximate a zero potential to which
the positive or negative change recorded
by the active input can be compared.

\lI

--

+30
0
'::;
-30 0
-60 :::!
- 90

if)

summed up in the P wave of the electro

cardiogram, gives rise to negative po
tentials on the chest above the atria and
positive potentials below. Recovery or
repolarization of the atria does not show
up in a body surface electrocardiogram,
because the potentials are small and are
hidden in the QRS complex.
From the intricate shape of the QRS
complex, on the other hand, one can
tell little about the flow of electrical
activity through the massive muscula
ture of the ventricles. Early investiga
tors (notably Frank N. Wilson of the
University of Michigan Medical School,
the father of modern electrocardiog
raphy, and Richard Ashman and Man
uel Gardberg of Louisiana State Univer
sity School of Medicine) made some
shrewd deductions, but it was clear that
the information furnished by body sur
face potentials would have to be sup
plemented by direct observation of elec
trical activity in the heart. To make such
observations we constructed a compound
electrode consisting of
fine tungsten
wires, the tips of which are exposed
along a shaft at one-millimeter intervals.
Inserted perpendicularly to the wall
of the heart in a laboratory animal, this
electrode makes it possible to record
potential changes acrOss the thickness
of the muscle as the wave of activity
sweeps by. A large number of insertions
are necessary to study the entire heart,
and the readings at each point are
recorded simultaneously along with a
"time reference" potential from a fixed
point in the heart. The activity is regis
tered by the multitip electrode in one
of two ways: by picking up the potential
between each of the tips and an electrode
at a distant point on the body, or by
recording the sharp "pip" that occurs
when the wave is midway between each
pair of tips along the length of the elec
trode. With as many as
insertions, and
therefore with readings from as many as
points, we have been able to chart
in three dimensions the multidirectional
pathway of the ventricular potential
changes. Our studies are corroborated by
the work of Demetr:o Sodi-Pallares at the
National Institute of Cardiology in Mex
ico and D. Durrer at the University of
Amsterdam.
Direct observation by this means
shows that the first regions in the ventri
cles to become excited are at the termi
nations of the conduction system, which
originates from the atrioventricular node.
The original excitation at the pacemaker
is relayed to these electrical conduits by
the wave of activity that sweeps through
the muscles of the atria. The right and

15

70

1,050

DIFFERENCE IN ELECTRIC CHARG E inside and outside cell membmne sets up a

potential. Resting cells

(left alld right)

have low internal sodium concentration

(hatchillg)

and high potassium concentration. Outside the opposite situation prevails. When cell be
comes active

(center),

sodium ions flow into it, reversing the charge distribution; the out

side is negative, the inside positive. Cell is now depolarized. Cmrent flows into it from
adjacent cells that have not been depolarized in this cycle and they rapidly become active.
The potential changes recorded from an active heart-muscle cell are shown at top. At rest
its interior is

90

millivolts negative with respect to the exterior. In active state it is

30

millivolts positive. It needs half a second to return to resting state. Broken line indicates
duration of same potential changes in a nerve fiber, which recovers in less than

.001

second.

134
1961 SCIENTIFIC AMERICAN, INC

r
\
J

++++++++++++ +
+
+
+
+
+++++++++++++
WAV E OF ACT IV ITY moves from left end of mllscle strip to

show a negative potential at left end of strip, a slight positIve

right end. Above, only the small block of tissue at far left is ill

potential at center and smaller positive potential at right edge.

active state (depolarized). Black sections of the three curves

The gray sections of the curves indicate changes that will follow.

++++++++
+

+
+

++++++++
WAV E MOV ES ON to center of strip. Strong negative potential is

potential is now approaching zero, while at the right a high posi

being recorded at lefthand electrode; in the center the positive

tive potential is being recorded as the active wave approaches.

-Y

------------ +++
+
+
+
+
+++
AS W AVE N EARS R IGHT

END, potential at left is zero. At

the right end of muscle strip registers a posItIve potential that

the center it is now negative, returning to zero. The electrode at

will return rapidly to zero as the wave passes out of the region.

135
1961 SCIENTIFIC AMERICAN, INC

left conduction bundles of the system

descend on each side of the interventric
ular septum and radiate from there over
the inner surface of the ventricles. The
units of the conduction system are so
small that their activity goes undetected
by recording electrodes on the surface

both sides. Because there is more active

tissue on the left side of the septum, the
left-to-right movement exceeds the right
to-left movement. This leads to the po
tential change recorded by the electro
cardiogram in the first part of the usually
triphasic QRS complex.

of the body. Ventricular excitation, as

shown by the multitip electrode, begins
in the inner surface of the heart on either
side of the septum about halfway be
tween the top and bottom of the cham
bers. The activity at first spreads inward
toward the center of the septum from

ATRIA AR E DEPOLARIZED by wave that moves downward and

to the left

right)

(colored arrows).

P waves produced by this activity

(far

are negative when recorded at left shoulder (1), positive

and negative from electrode at

and positive when recorded at

3.

These differences come about because the activity wave is moving

away from

1,

toward and away from

and toward the electrode at

J.

i
MOVEMENT OF DEPOLARIZING WAV E through the ventricles
'
produces a variety of curves of potential change

(colored arrows)

when recorded through electrodes located at four different spots

around the heart. The changes of potential are so complex that the

136
1961 SCIENTIFIC AMERICAN, INC

Soon after the earliest signs of ac

tivity in the septum, the fibers of the con
duction bundles distribute the excitation
over the inner surface of the lower and
middle portions of the free walls of the
ventricles. The branching of these fibers
is so extensive that the lining of the
ventricles in this region depolarizes al
most simultaneously. As a result the only
direction in which the wave of excitation
can move is outward, toward the outer
surface of the heart. This activity adds
up to a single wave directed downward
toward the tip, or "apex," of the heart
and generates the second phase of the
QRS complex.
The conduction bundles, which carry
impulses at a speed of one or two meters
per second, thin out toward the top, or
"base," of the ventricles and do not
reach the upper regions of the septum
or of the ventricular walls. These regions
are therefore the last to depolarize. They
are excited during the S wave of the
QRS complex by a wave conducted
through the muscle and moving upward
to the base at meter per second.

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.4

he three phases of ventricular ac-

T tivity-from right to left in the sep

tum, inside out in the walls, and toward

the base of the ventricles through the
walls and the septum-overlap in time
and succeed one another smoothly. In
the electrocardiogram the QRS shape
varies considerably, depending on where

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movement of the Q, R and S waves cannot
he analyzed by this standard technique.

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137
1961 SCIENTIFIC AMERICAN, INC

-20 -10

10

20

TIME (MILLISECONDS)

30

40

-V'iIl10 5 { 0
\

WAVE OF EXCITATION moves in the directions indicated by the arrows in vertical sec
tion of the ventricles

(middle).

Electrodes that measured the wave were inserted along the

axes of the arrows. At bottom is a horizontal section of the ventricles. Numbers and contour
lines indicate time at which the wave reached a specific region. Electrocardiogram is at top.

138
1961 SCIENTIFIC AMERICAN, INC

the electrodes are placed on the body.

Furthermore, the patterns recorded
from the same sites on two normal people
may vary, due to such anatomical pecu
liarities as differences in the distribution
of the conduction tissue, the thickness of
the heart walls and the position of the
heart in the chest.
During the Rrst p hase of ventricular
depolarization as it is seen by the electro
cardiograph, the wave moves approxi
mately toward the right arm, and an
electrode placed there records a posi
tive potential. Because the left arm is
virtually perpendicular to the direction
of travel, an electrode there records little
activity. The electrode on the right side
of the chest, like the one on the right
arm, records approaching activity; the
electrode on the left side of the chest,
receding activity. In the second phase
the wave can be considered to be mov
ing toward the left leg, since this is the
sum of all the inside-out activity in the
walls. The electrode on the right arm
records a large amount of recedmg ac
tivity and that on the left arm a lesser
amount. The electrode on the right side
of the chest also records receding activ
ity, while that on the left indicates a
large amount of approaching activity.
With the wave moving toward the head
in the Rnal phase of ventricular excita
tion, the electrodes on the arms record
approaching activity and the electrodes
on the chest record receding activity.
The T wave, which corresponds to
repolarization, or electrical recovery, of
the ventricular muscles, still presents a
puzzle. In a human electrocardiogram
this wave has the same polarity as the
QRS complex in recordings from many
body surface points. It should, of course,
be opposite in sign, because the poten
tial changes across the boundary between
resting and active tissue are opposite to
each other during depolarization and
repolarization. This suggests that the
pathway of repolarization in the human
heart may take a different-perhaps op
posite-direction from depolarization. Al
ternatively, one might conclude that
repolarization does not spread from cell
to cell as a result of current flow. Instead,
the sequence in which the cells repolar
ize may be determined by metabolic
events and have only a statistical direc
tion. Experimental studies with animals
cannot yet answer this question because
no animal has a T wave like that in man.
Certain forms of cardiac abnormality
are simply and unequivocally demon
strated by the electrocardiogram. For
example, the P wave and the QRS com
plex normally occur at the same frequen
cy, and the latter follows the former at a

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The head of the periscope breaks the surface, star
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In
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139
1961 SCIENTIFIC AMERICAN, INC

constant interval. In many disorders of

cardiac rhythm this relationship is al
tered either by loss of the one-to-one
relationship of the waves or by an abnor
mality of the interval between the waves.
Another type of defect, referred to as a
conduction defect, can arise from dam
age to the specialized conduction tissue
of the atrioventricular node or conduct
ing bundles. If the defect blocks the ac
tion of the conduction bundle leading to
the right ventricle, the depolarization of
the left ventricle will appear normal, but

since the wave of activity can reach the

right ventricle only by traveling slowly
across the interventricular septum, it
will show up late. The QRS complex is
prolonged and characterized by move
ment of activity from left to right in its
terminal portion.
In coronary thrombosis, the most com
mon form of "heart attack," the electro
cardiogram yields direct information
about the extent of damage immediately
following the episode. Normally the elec
trocardiogram does not show potentials

except during the

QRS and T waves;
in the intervals between these electrical
events the potential is zero. But when
segments of ventricular muscle are de
prived of oxygen by loss of blood supply,
as occurs in coronary thrombosis, the pen
does not return to the pre-QRS base line.
This potential is called "current of in
jury" and arises from the fact that the
cells are partly depolarized by oxygen
deprivation.
Later, when the injured muscle has
been replaced by electrically inactive

P,

NORMAL
A

A
B

20

60
40
TIME (MILLISECONDS)

100

80

DEPOLARIZATION IN V ENTRICLES is shown in four cross sec

30

milliseconds, as seen in the electrocardiogram at left. Crisscross

is at the atrioventricu

lines in the sections at zero milliseconds show locations of elec

lar valve. The complete cycle goes smoothly to completion in about

trodes implanted for author's observations. The color in the sec-

tions of normal dog heart. Top of section

20

80
60
40
TIME (MIlliSECONDS)

100

DAMAGE TO HEART can change pattern of depolarization. In

by spreading slowly across septum as depolarization of one cell

this case the left conducting bundle has been cut. Activity begins

triggers activity in the next. The QRS complex is prolonged and

around right cavity of dog heart, but can reach left side only

characterized by delay in the movement of activity. The electro

140
1961 SCIENTIFIC AMERICAN, INC

scar ttssue (a condition termed chronic

myocardial infarction) , the potentials
normally contributed by the destroyed
muscle will be absent from the electro
cardiogram. The interpretation of the
electrocardiogram in this phase of the
affliction is more complex, but useful cri
teria of damage have been accumulated
through experience with many patients.
This clinical experience is gradually be
ing supplemented by understanding of
the details of function that underlie the
normal electrocardiogram.

CUT YOUR
INDICATOR LIGHT
COSTS!
Use GE Glow Lamps

with resistor attached

tions at 5, 12,
resents

the

18

and 25 milliseconds rep

spread

of

depolarization

in

ventricles as the QRS complex takes place.

You may be paying more than you should for your indicator
lights if you're using anything but General Electric glow lamps
with the resistor attached. The popular NE-2H and NE-2E,
for instance, in large quantities, cost less than seven cents
apiece including the resistor.
Low cost isn't the only reason why G-E glow lamps make
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Other big advantages of G-E glow lamps are their rugged
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word to the wise: if you're designing appliances, business

machines or military hardware, get the story on General
Electric glow lamps as indicators. There are over 60. Write:
General Electric Co., Miniature Lamp Dept.
Park, Cleveland

12, Ohio.

M-145,

Nela

Progress Is Ovr Mos! Imporfanf Prot/vel

cardiogram plainly reflects this abnormal
condition. The cycle lakes
instead of

30

90

milliseconds

milliseconds for completion.

GENERAL

fj ELECTRIC
141

1961 SCIENTIFIC AMERICAN, INC

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trol with the UNIVAC Solid-State Computer.
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Just this: this is the first time a low-priced, gen
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over

300

How does Univac numerical control work? With

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1961 SCIENTIFIC AMERICAN, INC

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brief you on Univac numerical control-for your own

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DIVISION

1961 SCIENTIFIC AMERICAN, INC

OF SPERRY RAND

CORPORATION