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tion becomes feasible. was considered to be unfolded into the plane of the
interatrial septum as shown in figure lB. In this
METHODS AND MATERIALS figure, point S represents the site of impulse forma-
In the analysis to be presented, the atria were tion in relation to the interatrial septum. Since the
considered as two surfaces. One of these consisted distance which the activation wave must travel
of those portions of the free walls of the atria which from the point of origin to some portions of the left
were visible on the external surface of the heart (free atrium is shorter via the septum than through the
external atrial walls). The remainder of the atria, free atrial walls, the point S was also located in
which included the interatrial septum and those relation to the external wall of the left atrium as
portions of the free walls which were not visible on shown in figure 1C. In this figure the free external
the external surface of the heart (free internal atrial atrial walls have been "unfolded" onto a plane
walls), were considered to form the second surface. surface and the septum has been divided to show the
Activation was assumed to begin on the free ex- apparent location (S) of the site of origin of impulses
ternal atrial wall at the anterior margin of the junc- which pass into the free wall of the left atrium from
tion of the superior vena cava and the free external the septum. The point F represents the site of origin
wall of the right atrium. This point represents the of impulses which pass directly into the free wall of
approximate location of the superior portion or the right atrium.
head of the sinoatrial node as described by Keith The atrial outlines used in this study were ob-
and Flack.1 It was further assumed that the wave tained by dissecting human atria in the following
spread in all directions at uniform speed. This fashion. The unopened atria were filled with cotton
introduced through the atrioventricular valves from
From the Department of Medicine, Tulane Uni- the opened ventricular cavities. The approximate
versity School of Medicine, Charity Hospital of portions of the free external walls which faced
Louisiana at New Orleans, and the Department of anteriorly, laterally and posteriorly with the heart
Electrical Engineering, Tulane University of Louisi- approximately in the anatomic position were de-
ana, New Orleans, La. termined. A rough outline of the margins of the in-
Aided by a grant from the Public Health Service tact atria in their approximate anatomic position
(H-143) and the Mrs. E. J. Caire Fund for Research was obtained by wrapping a strip of tracing paper
in Heart Disease. around the atria and tracing their outline. Vertical
97 Circulation, Volume XI, January, 1955
98 ANALYSIS OF ACTIVATION IN HUMAN ATRIA
Free internal wall B.
A. Sup. veno / L. atrfum
B- s with the frontal plane and with the sagittal plane is
cava s^ illustrated in figure 3. Figure 3A shows a coplanar
outline of the unrolled free external atrial walls and
its division into anterior, posterior and lateral
portions, and 3B shows the spatial orientation these
SF portions of the atria were considered to have in this
study. Figure 3C shows the coplanar outline of the
septum and the free internal atrial walls, and figure
3D shows the spatial orientation these portions were
Inf. vena cava
assumed to have.
The spread of the activation wave was represented
on the unfolded surfaces described as a series of
concentric arcs drawn about the site of impulse
formation. Since the site of impulse formation was
represented by two points in relation to the free
external atrial walls a set of arcs was drawn about
each of these points.
In each of the sections described, the length of
FroIG. 1. The relations of the pacemaker in the sino- the chord of an arc defined by the limits of the
atrial node are diagrammatically illustrated. A section depicting a given moment in the excitation
shows the unopened atria with the pacemaker repre- process was taken as the magnitude of an electric
sented by the point SF. The septum and the internal vector acting at that moment. This vector was
surface of the free wall of the left atrium are shown in assigned a direction perpendicular to the chord
black. The internal surface of the free wall of the right
atrium is not shown. B shows a section of free right
atrial wall containing the pacemaker (S) unfolded A.
into the plane of the septum. C shows a coplanar out-
line of the external surface of the free atrial walls
with the pacemaker shown at F and at S.
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4A shows the outline of the unrolled free external cd and perpendicular to that chord. In C, vectors
walls of the atria of a normal heart and 4B shows V' and V2 have been added to give vector V3. Vector
the septum and the free internal atrial walls. The V4 has been made equal in length to the chord ef and
directed perpendicular to that chord. In D, vectors
A. B. V3 and V4 are added to give V5 which is the effect in
the left sagittal plane of activation at the stage
represented by the arcs labeled 3 and 3'.
site of impulse formation is represented by points
F and S. As discussed, both F and S represent a
single site of impulse formation, but F shows this
site in its normal relation to the free wall of the
Rt. Ant. L Ant. right atrium while S represents the apparent point
of origin of impulses which entered the free atrial
C.
walls from the septum. The atria have been divided
Free internol wall into portions parallel to the frontal and sagittal
L. atrium
R. atrium
planes (figs. 4A and B).
In figures 4A and 4B, a series of concentric arcs
about the points representing the site of impulse
formation are shown. Each of the arcs about a
single point represents the activation process at a
given moment. The arcs were numbered consecu-
tively; those which occurred simultaneously were
FIG. 3. Division of the atria into sections parallel designated by the same number, and those as-
with the frontal and sagittal planes is illustrated. sociated with B were primed. The derivation of
The coplanar outline of the free external walls of a vectors from two simultaneous arcs (numbers 3
normal specimen is shown in A, and its assumed and 3') in the lateral portions of the atria is il-
orientation is shown in B. The septum and free in- lustrated. The vector V' was assigned a magnitude
ternal walls are shown in C, and their assumed spatial equal to the length of the chord CD and a direction
orientation in D. perpendicular to that chord. Similarly, vector V2
100 ANALYSIS OF ACTIVATION IN HUMAN ATRIA
was assigned a magnitude equal to the length of form of the P waves recorded electrocardiographi-
the chord EF and a direction perpendicular to that cally with those constructed as outlined. The spatial
chord. Vectors V1 and V2 were then oriented as vectorcardiogram of one patient recorded with the
they would have been in the unopened atria and equilateral tetrahedron reference system was avail-
added vectorially to give V3 as shown in figure able for comparison with the frontal and sagittal
4C. A simultaneous vector, V4, was derived from plane projections of the constructed Psti-loop.
arc number 3' as it appeared on the interatrial
septum. Vectors V3 and V4 were added vectorially RESULTS
to give vector V5 which was the resultant in the
sagittal plane of that stage in the activation process Normal Atria. In the two normal atrial
represented by arcs 3 and 3'. The vector repre- specimens analyzed, the relative amplitudes
senting the manifestation in the frontal plane of of the derived P waves were similar to those
activation at this moment was obtained by carrying of the P waves in the standard leads of the
out a similar procedure using those portions of the
same arcs which were within the anterior and pos- electrocardiogram recorded before the patient's
terior sections of the atria. death. The general contour of the derived
When this procedure was carried out with con- waves was remarkably similar to that of the
centric arcs placed 1 cm. apart, a series of vectors recorded P waves although some differences
representing the effect of activation in the frontal in form were present (fig. 5).
and sagittal planes at several sequential intervals Recorded Psi-loops of these patients were
was obtained. Joining the termini of these vectors
in their proper time sequence gave figures analogous not available for comparison with the derived
to the frontal and sagittal plane projections of the projections; however, the configuration and
Psi-loop. orientation of the derived loops were similar
P waves were derived from the loops in the con- to normal PsiS-loops recorded in this labora-
ventional fashion for comparison with those actually
recorded in the standard leads. The time scale was tory.
arbitrarily chosen to conform to average normal The coplanar outline of one normal atrial
P-wave time-amplitude relationships. specimen was shown in figure 4. Figure 5A
Seven human atria were employed in this analy- shows the frontal and left sagittal plane pro-
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sis. Two of the specimens were atria of normal jections of the Psi-loop derived from this
size, two were from patients with rheumatic dis- specimen. The derived P waves are shown in
ease of the mitral valve and had left atrial enlarge-
ment, and three showed right atrial enlargement, figure 5B, and the electrocardiographically re-
two resulting from cor pulmonale and one from corded P waves are shown in figure 5C. The
congenital pulmonary stenosis. Electrocardiograms general form and the relative amplitudes of the
showing a normal sinus mechanism were available derived waves may be seen to be similar to
on all subjects making it possible to compare the those in the recorded electrocardiogram.
Left Atrial Enlargement. In the two speci-
mens with left atrial enlargement studied, the
~ ~ _ :-
'!
r^: i: ^
relative amplitudes and general contour of the
derived waves were similar to those of the
'*\ I· ' i~:-sl a ti P waves in the electrocardiogram recorded
prior to the patient's death. Both the derived
Hr
B~~
-I /v
lr ^u-~it waves and the electrocardiographically re-
corded P waves had a longer duration than
those of the subjects with atria of normal size.
Lft
sagittolI m li1----
Elr In each instance the notched wave which is
usually associated with left atrial enlargement
: 1* , * u1 111ni occurred in both the derived and the actual
P waves in lead I.
FIG. 5. A shows the frontal and sagittal plane pro- Since the characteristics of the Psti-loop
jections of the PsE-loops derived from the normalP in left atrial enlargement are not established,
atrial specimen shown in figure 4. The derived
waves are shown in B, and the P waves of the electro- comparison with actual recorded loops could
cardiogram recorded before the patient's death are not be made. The frontal plane projections of
shown in C. the two derived Psti-loops were located in the
J. A. ABILDSKOV, J. A. CRONVITCH, AND G. E. BURCH 101
fifth and sixth sextants of a triaxial reference $
system. The left sagittal plane projection of
one was located in the fifth and sixth sextants
while the other was confined to the fifth sextant
of a triaxial reference system applied to that
plane with the =-180 degree axis located an-
teriorly. Both frontal plane projections and one
sagittal plane projection had figure-eight con-
figurations, while the left sagittal plane pro-
jection of the other was a simple loop inscribed
in a clockwise direction. Characteristic of both
frontal plane projections was a rapid movement
of the loop to the right and then to the left re-
sulting in a notching of the derived wave in
lead I. FIG. 6. The coplanar outline of a specimen showing
The coplanar outline of one of the specimens left atrial enlargement is shown in A. The scale of
showing left atrial enlargement is shown in this and the following illustration is indicated by the
concentric arcs which were placed 1 cm. apart. The
figure 6A. Figure 6B shows the frontal and frontal and sagittal projections of the PsE-loops
left sagittal plane projections of the PsiS-loop derived from this specimen are shown in B, and the
derived from this specimen. The derived P derived P waves and the P waves in the electrocardio-
waves are shown in figure 6C, and the electro- gram are shown in C and D.
cardiogram recorded prior to the patient's
death is shown in figure 6D. The general form derived Psti-loops were located in the fifth
and relative amplitudes of the derived and re- and sixth sextants of the triaxial reference
corded P waves are similar. system. Two of the sagittal plane projections
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Right Atrial Enlargement. Three atrial speci- were located entirely within the fifth sextant,
mens showing enlargement of the right atrium while one was within the fifth and sixth sextants
were studied. When right atrial enlargement of a triaxial reference system applied to that
was present, a greater amount of atrial muscula- plane. All frontal plane projections were in-
ture was found to lie parallel to the frontal scribed in a clockwise direction, with two of the
plane than in the atria of normal size and those records so inscribed that the line crossed itself
with left atrial enlargement. In the method near the distal tip of the loop. Two of the
employed in this study the increased amount sagittal plane projections were inscribed in a
of muscle so located resulted in vector forces clockwise direction, and one formed a figure-
of increased magnitude in the frontal plane. eight.
This in turn was reflected in the derived P The coplanar outline of a specimen showing
waves as increased amplitude in one or more massive enlargement of the right atrium is
leads. Such increased amplitude of the P waves shown in figure 7A, and the frontal and left
is consistent with the clinical observation of sagittal plane projections of the Psti-loop de-
high peaked P waves in the presence of right rived from this specimen are shown in figure
atrial enlargement. As was true of the normal 7B. The frontal and left sagittal plane projec-
atria, and those showing left atrial enlarge- tions of the spatial vectorcardiogram recorded
ment, the general contour and the relative prior to this patient's death with the equilateral
amplitudes of the derived waves were similar tetrahedral reference system are shown in
to those of the P waves in the electrocardio- figure 7C. The orientation, general form and
grams recorded prior to death. Both the de- direction of inscription of the derived projec-
rived and recorded waves of the three subjects tions of the Psi-loop were similar to those of
studied were high and peaked and of greatest the Psti-loop actually recorded. The P waves
amplitude in lead II. derived for the standard leads are shown in
The frontal plane projections of the three figure 7D, and the P waves recorded simul-
102 ANALYSIS OF ACTIVATION IN HUMAN ATRIA
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FIG. 7. The coplanar outline of a specimen showing massive enlargement of the right atrium ;s
shown in A. Frontal and sagittal projections of the PsE-loops derived from this specimen are shown
in B, and these projections of the PsE-loop recorded with the equilateral tetrahedron reference sys-
tem are shown in C. P waves plotted from the frontal plane projections of the derived PsE-loops are
shown in D, and the P waves recorded in simultaneous pairs are shown in E.
J. A. ABILDSKOV, J. A. CRONVITCH, AND G. E. BURCH 103
taneously in pairs prior to the patient's death from the site of impulse formation. This order
are shown in figure 7E. The general form and of excitation is supported by the studies of
relative amplitudes of the derived waves may Lewis5 and of Prinzmetal and co-workers.6
be seen to be similar to those of the recorded The observations of Eyster and Meek,7 how-
P waves. It may also be noted that the peak ever, led them to conclude that even though
of the P wave in lead III of both the recorded no specialized conducting tissues exist in the
electrocardiogram and the derived waves occurs atria, impulses spread preferentially over cer-
later than the peak in leads I and II. The tem- tain paths to reach the atrioventricular node.
poral relationships of various portions of the The recent studies of Puech and associates3
derived P waves agree remarkably well with on auricular activation in the dog's heart
those of the recorded P waves. although showing a virtually constant con-
duction speed in the right atrial body and ap-
DISCUSSION
pendage found it always higher in the region
Because of the limited state of knowledge of the taenia terminalis. Results of the present
several assumptions which are known to be study appear to support the view that activa-
only partially correct have been made in this tion of the atria proceeded in a regular radial
analysis to facilitate the calculations. For ex- fashion, since the form of the derived Psi-
ample, the treatment of the atria as plane sur- loops and P waves closely resembled those
faces is an oversimplification. The free walls actually recorded. It is possible, however, that
of the atria visible on the external surface of preferential conduction paths to the A-V node
the heart are actually more nearly oriented in operated without significant influence on the
the form of a crescent with no sharp boundaries form of the Psfi-loop and P waves while the
between anterior, lateral and posterior surfaces. bulk of the atrial musculature was activated
Likewise, the orientation of the septum is more in a radial fashion.
complex and more variable than has been as- In this analysis the site of impulse formation
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sumed in this study. Thus some portions of the was assumed to lie at the anterior margin of
atria which were considered to influence only the junction of the superior vena cava and the
the form of the frontal plane projection also right atrium. This point corresponds to the
contribute to the form of the sagittal plane superior portion of the collection of sinoatrial
projection of the Psi-loop, and some portions nodal tissue originally described by Keith and
which were considered to influence only the Flack1 and represents the approximate locus
sagittal plane projection influence also the form which Lewis2 and others3' 7 concluded was
of the frontal plane projection. Furthermore the pacemaker of the normal heart. Since the
the superior and inferior surfaces of the atria form and orientation of the Psti-loops and the
have not been considered in this study. Another form of the P waves derived in this study were
source of error was the failure to take into ac- dependent on the site of impulse formation,
count all the physiologic openings in the atria. these results tend to support the view that
Simultaneous depolarization and repolariza- impulses originated in the head of the sinoatrial
tion in the atria may have further limited this node.
analysis. MacLeod's observations4 indicated In spite of the limitations, the analysis pre-
that repolarization in the frog atria began be- sented appears to have value in understanding
fore depolarization was completed and this the form of normal and abnormal Psti-loops
may be true for man as well. Modifications of and P waves. In each instance the derived P
the method reported would be suitable for waves were similar in contour and in relative
studies relevant to this problem. It was not amplitude to those actually recorded before
possible to assess the importance of the errors the subject's death, and in the case of the one
introduced by the simplifying assumptions em- subject whose spatial vectorcardiogram was
ployed in these studies. available the PsiS-loop was similar to that
The excitation wave was considered to spread which was calculated. In general, the derived
through the atria in a uniform radial fashion P waves had a more complex form than was
104 ANALYSIS OF ACTIVATION IN HUMAN ATRIA
apparent in the P waves of the electrocardio- one patient recorded prior to death was also
gram. This, however, may have been at least similar in general form, orientation and direc-
partially due to the lack of detail in the P wave tion of inscription to the Psi-loop derived
recorded electrocardiographically. Studies re- from the atrial outline.
ported by ILangne'r, in which the cathode ray 3. Left atrial enlargement resulted in de-
oscilloscope with an expanded time scale and rived P waves of greater duration than those
amplifiers with high frequency response were derived from normal atria. Rapid changes in
used, demonstrated details in the configuration the direction of inscription of the frontal plane
of the P wave which were not apparent in those projections of the derived PsE-loops were as-
recorded with the conventional electrocardio- sociated with notching of the derived P wave
graph. in lead I. These characteristics of the derived
The analysis outlined accounted for some Psi-loops and P waves were clearly the result
of the differences in normal P waves and Psf- of the size, contour and orientation of the atria.
loops and those occurring with atrial enlarge- 4. Enlargement of the right atrium resulted
ment. Right atrial enlargement resulted in in a greater mass of atrial musculature oriented
frontal plane vectors obtained by the method parallel to the frontal plane. This produced
outlined which were of greater magnitude than vectors of increased magnitude in the frontal
those derived from normal atria. This was a plane projection of the "derived" PsE-loops
consecluence of the greater mass of atrial mus- and was reflected by high, peaked derived P
culature located anteriorly which produced waves in the standard leads.
high peaked P waves in some of the standard SUMMARIO IN INTERLINGUA
leads of the electrocardiogram. The duration
of the derived P waves in atrial enlargement 1. Septe atrios human esseva dissecate de
was longer than in normal atria. These charac- maniera a permitter le executionl de un delinea-
teristics of form and duration are consistent tion coplanar de lor contorno. Per supponer
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with those observed in the electrocardiograms que le sito normal del formation del impulso se
of patients with right atrial enlargement. The trova al capite del nodo sino-atrial e que le
form and the orientation of the atria in which unda de activation se expande radialmente ab
the left atrium was enlarged gave rise to cal- iste puncto, nos poteva obtener "derivate"
culated Psi-loops of increased duration and of projectiones planar del spiras PsI; e del ullda
such a form that prominent notching of the P in derivationes standard.
derived P waves in one of the standard leads 2. Le derivate undas P se monstrava simile
was present. Such P waves are known to occur in amplitude relative e contorno general al
commonly with left atrial enlargement. undas P registrate ante le morte. Un vecto-
SUMMARY
cardiogramma spatial registrate ante le morte
in le caso de un del patientes esseva etiam
1. Seven human atria were dissected in a simile in forma general, orientation e direction
manner which allowed a coplanar outline of de description al spira PsE derivate ab Ie ccn-
their contour to be made. By assuming that torno atrial.
the normal site of impulse formation was lo- 3. Allargamentos sinistroatrial resultava in
cated at the head of the sinoatrial node and durationes del derivrate unda P plus ext.endite
that the activation wave spread ill a simple que in le case de atrios normal. Rapide cambia-
radial fashionii from this point, it was possible mentos in le direction del inscription de pro-
to obtain "derived" planar projections of the jectiones inle plano frontal de derivate spiras
PsE-loops and standard lead P waves. PsE esseva associate con indentationes in le
2. The derived P waves were found to be derivate unda PI del derivation I. Iste charac-
similar in relative amplitudes and general con- teristicas del derivate spiras PsE e del undas P
tour to those recorded electrocardiographically esseva clarmente le resultato del dimensiones,
prior to death. A spatial vectorcardiogram of del contorno, e del orientatioln del atrios.
J. A. ABILDSKOV, J. A. CRONVITCH, AND G. E. BURCH 105
4. Allargamentos del atrio dextere resultava AND CISNEROS F.: Normal auricular activation
in uni augmento del massa de musculatura in the dog's heart. Am. Heart J. 47: 174, 1954.
4 cMACLEOD, A. G.: The electrogram of cardiac
atrial que esseva orientate parallelmente al muscle. An analysis which explains the regres-
planio frontal. Isto produceva vectores de aug- sion of T deflection. Am. Heart J. 15: 165, 1938.
mentate magnitude in le projection in le piano 5
LEWIS, T.: The Mechanism and Graphic Registra-
frontal de "derivate" spiras Psfi e esseva re- tion of the Heart Beat, ed. 3. London, Shaw,
flectite per derivate undas P a alte piccos in le 1925.
derivationes standard.
6
PRINZMETAL, 1M., CORDAY, E., BRILL, I. C.,
OBLATTH, R. W., AND KRUGER, H. E.: The
REFERENCES Auricular Arrythmias. Springfield, Ill., Charles
KEITH, A.,
' AND FLACK, MI.: The form and nature C Thomas, 1952.
of the muscular connections between the prim- 7
EYSTER, J. A., AND M1EEK, W. J.: Experiments on
ary divisions of the vertebrate heart. J. Anat. the origin and propagation of the impulse in
& Physiol. 41: 172, 1907. the heart. Heart 5: 119, 1913-14.
2 LEWIS, T.: The site of origin of the rmammsalian LANGNER, P. H.: The value of high fidelity elec-
heart beat; the pacemaker in the dog. Heart trocardiography using the cathode ray oscillo-
2: 147, 1910. graph and an expanded time scale. Circulation
3
PUECH, P., ESCLAVISSAT, Mi., SODI-PALLARES, D., 5: 249, 1954.
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