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Thorax (1957), 12, 304.

THE INTERVENTRICULAR SEPTUM

BY
E. W. T. MORRIS
From the Anatomy Department, St. Thomas's Hospital Medical School, Londoni

(RECEIVED FOR PUBLICATION JULY 26, 1957)

It is difficult to find in the literature a clear and between the tips of its two horns where the
concise account of the development and form of boundary is formed by the fused atrioventricular
the interventricular septum. Moreover, some of cushion (A, in Fig. 4). This septum does not lie
the accounts in the clinical literature are at vari- in one plane and the main part of its free border
ance with that generally accepted by embryo- forms a spiral (Figs. 4 and 5).
logists. For this reason and in view of the recent (2) While the muscular part is forming. changes
technical advances in the surgery of the heart, it are taking place in the relative positions of the
seems opportune to describe the development and bulbus cordis and the ventricles. Earlier the heart
anatomy of the interventricular septum and to tube is flexed at the bulboventricular junction so
correlate this knowledge as far as possible with that the bulbus cordis comes to lie ventrally and
the sites of interventricular septal defects. to the right of the ventricle (Fig. 2). Their con-
At an early stage the heart consists of the sinus tiguous walls form a septum-the bulboven-
venosus, the common atrium, the common ven- tricular septum-around the lower free border
tricle, and the bulbus cordis, serially arranged in of which the two cavities communicate (see Figs.

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that order from the venous to the arterial end 1, 2, and 3). This septum intervenes between the
(Figs. 1 and 2). distal part of the bulbus cordis and the atrio-
The formation of the interventricular septum is ventricular opening (Fig. 3). It must disappear
a complicated process involving the partitioning before the interventricular septum can be com-
of the common ventricle into a right and left, and pleted.
the separation of the distal part of the bulbus The absorption of the bulboventricular sep-
cordis into pulmonary and aortic outflow chan- tum takes places whiel>the muscular septum is
nels in continuity with the ventricles, the proxi- forming, and, as a result3nd of unequal growth
mal portion of the bulbus cordis forming the rates in the different parts, the cavity of the upper
infundibulum of the right ventricle. part of the bulbus cordis comes to lie astride the
The process begins at about 5 mm. crown- middle and dorsal parts of the upper free border
rump length (35th day) and normally ends about of the muscular septum (Fig. 4). The next step
17 mm. crown-rump length (49th day). has meanwhile begun; it is the division of the
The septum is formed from the following struc- distal part of the bulbus cordis by two ridges, the
tures, which begin their contribution in this right and left bulbar ridges, which grow from its
order: (1) the muscular wall of the common walls and ultimately fuse. These ridges start dis-
ventricle, (2) the bulbar ridges, and (3) the dor- tally and grow proximally (caudally). One grows
sal atrioventricular cushion. down the right side of the dorsal wall of the bulb
(1) The muscular contribution starts as a ridge to the right end of the ventral component of the
on the dorsal wall (Fig. 3) and extends on to the fused atrioventricular cushions opposite the
ventral wall of the common ventricle. It is then attachment of the dorsal horn of the muscular
crescentic in form and its dorsal horn reaches the septum to the dorsal component. In its progress
right end of the dorsal atrioventricular cushion. this ridge grows over and obliterates the ventral
The ventral horn approaches the ventral atrio- part of the right atrioventricular canal (Fig. 4).
ventricular cushion near its centre (Fig. 4). While The other ridge grows down the left side of the
this septum is forming the central parts of the ventral wall of the bulbus cordis and approaches
atrioventricular cushions fuse (11 mm. C.R.L.). and fuses with the muscular septum on its right
The ventricles now communicate by a foramen, side a little distance along its margin from its
the boundary of which is formed mainly by the ventral end (Figs. 4 and 5). The ventral end of
free border of the muscular septum, except the muscular septum lies to the left of the left
 V~
~
~
~
i

9_
~~~~~~~~~~~~~~~~~~~~~~BVS
~~
{gV.
)2~
THE INTER VENTRICULAR SEPTUM

A.T

bulbar ridge. When the interventricular septum

is completed the part of the original interven-
tricular foramen, which is bounded by the ventral
horn of the muscular septum and. the fused atrio-
ventricular cushions between the horn tips, also s I
lies to the left of the completed interventricular '*.i
septum and so in the outflow channel of the left
ventricle, and the ridge formed by the ventral
horn is subsequently absorbed leaving a smooth
wall. The interatrial septum joins the atrial aspect
of the fused atrioventricular cushions about their
centre. As a result the fused atrioventricular
cushions between the attachment of the inter-
atrial septum and the dorsal horn of the muscu-
lar septum intervene between the outflow part of
the left ventricle and the right atrium (Fig. 9).
This becomes reorientated and finally lies in the
plane of the interventricular septum. In the final
form of the heart it becomes the atrioventricular
part of the pars membranacea septi (Fig. 7C).
At this stage in the formation of the interven-
"-

'
A.T.
B.C.
A.V.C.
B.V.S.

/;
,
\q~ ~ B.C
l
sinus venosus. A.T., common atrium.
A.V.C., atrioventricular canal.
common ventricle. B.C., bulbus cordis.
B.V.S., bulboventricular septum.

FIG. 2.-Ventral view of heart in

A
>as--in Fig. I.

commencing
andoVcar
mn sbaig.
Fig. 1.

V
305
FIG. 1.-Left lateral view of foetal heart. S.V.,
V.,

Key

FIG. 3.-Model of posterior part

of the bulbus cordis and
common ventricle. M.S.,
muscular inter-
ventricular septum. D.C.
ventriCularsepu.dv
doralauhind.ventaC.
sbendocardialcushions. Re-
Thorax: first published as 10.1136/thx.12.4.304 on 1 December 1957. Downloaded from http://thorax.bmj.com/ on September 19, 2023 by guest. Protected by
T ~~~~~~~~~~~~~~~~~~~~~~~

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tricular septum the distal bulbus cordis is divided

by a septum with a lower free border which forms V.C.
the upper boundary of a deficiency in the inter-
ventricular septum, the lower boundary of which A-V_ C

is formed by the upper free border of the muscu- D.C.

lar part (Fig. 6). Through this septal defect the
two ventricles communicate. V.
(3) This aperture is closed and the septum thus
completed by tissue derived from the dorsal atrio- M.S.
ventricular cushion in the neighbourhood of the
attachment of the right bulbar ridge and the
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306 E. W. T. MORRIS
FIG. 4.-Same view as in Fig. 3 at a later stage. A, fused endo-
cardial cushions. D.M.S.. dorsal part of the muscular
interventricular septum. V.M.S., ventral part of the
muscular interventricular septum. R.B.R., right bulbar
ridge growing down the wall of the B.C. and finally
obliterating the ventral part of the right atrioventricular
canal. L.B.R., left bulbar ridge (cut edge).

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,
L.B.R.

V.M.S.

D.M.S.

FIG. 5.-The anterior part of the same model as Fig. 4. Key as in

Fig. 4.
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THE INTER VENTRICULAR SEPTUM 307
A.O. FIG. 6.-Model of a foetal heart at a later stage
than Figs. 4 and 5. The bulk of the wall
of the future right ventricle has been
removed. P.T., pulmonary trunk. A.O.,
aorta. Y, site of origin of tissue from
dorsal endocardial cushion which com-
pletes the interventricular septum. Other
symbols as in Fig. 4.

A needle passed through the sep-

tum in the commissure between
the septal cusps of the aortic
valve emerges into the right ven-
tricle at the commissure between
the corresponding cusps in the
pulmonary valve (Figs. 7 and 8).
A. PART FORMED FROM DORSAL
- ATRIOVENTRICULAR C U S H I O N.-
This forms the interventricular
part of the pars membranacea
A.V.C.(R.) septi. It lies anterior to the atrio-
ventricular part of the pars mem-
-M.S. branacea septi and the whole pars
membranacea septi lies below and

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behind the bulbar septum and
above the middle of the upper
border of the muscular septum
(Figs. 7 and 8). As seen from
the left ventricle, the pars mem-
branacea septi lies below the com-
missure between right septal and
dorsal end of the muscular septum (Fig. 6). This non-septal cusps of the aortic valve and extends
tissue grows along and becomes fused with the forwards below the adjacent half of the right
edges of the aperture from dorsal to ventral. septal cusp (Fig. 7).
The anterior part of the attached border of the
THE ANATONIICAL SITUATION OF THE EMBRYOLOGI- septal cusp of the mitral valve crosses the pars
CAL COMPONENTS IN THE FINAL FORM OF THE membranacea septi between the atrioventricular
INTERVENTRICULAR SEPTUM and interventricular parts (Figs. 7 and 8).
THE MUSCULAR PART.-This forms the bulk of
the septum. OTHER FEATURES OF THE RIGHT VENTRICULAR
SURFACE OF THE INTERVENTRICULAR SEPTUM
THE BULBAR SEPTUM.-This forms the division
between the outflow channels and is continuous CRISTA SUPRAVENTRICULARIS.-From the point
below with the anterior part of the muscular sep- of view of classifying septal defects the impor-
tum. As seen from the left ventricle it forms the tant structure is the crista supraventricularis. This
part of the interventricular septum below the com- is a smooth ridge of myocardium in the anterior
missure of the two septal (coronary) cusps of the wall of the right ventricle lying parallel and close
aortic semilunar valve and the adjacent parts of to the attachment of the large anterior cusp of the
these cusps. This aspect of this part of the sep- tricuspid valve. It enlarges as it travels towards
tum contains no ventricular muscle. From the the left and ends by fusing with the interventricu-
right ventricular aspect this part is seen to form lar septum between the part formed from the bul-
the upper anterior part of the interventricular bar ridges and the interventricular part of the pars
septum which lies below the corresponding com- membranacea septi.
missure in the pulmonary valve. On this side, CONUS MUSCLE.-Arising from the septum just
the septum is muscular up to the semilunar valve. below the attachment of the crista supraventricu-
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308 E. W. T. MORRIS

_-._.~
; \.
.e'SEr " \~
i o-ni

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-e

laris are a group of papillary muscles to which are through the trigonum fibrosum dexter and thence
attached the chordae tendinae of the adjacent into the lower border of the atrioventricular sep-
parts of the anterior and septal cusps. One of tum, which lies just anterior to the trigone. It
these is usually larger than the rest, and this is passes from the atrium into the ventricle at the
then called the conus muscle. junction of the atrioventricular septum and inter-
ventricular part of the pars membranacea septi
CONDUCTING SYSTEM and runs in the latter at its fusion with the muscu-
ATRIOVENTRICULAR BUNDLE.-Arising from the lar septum or on either side of the upper border
atrioventricular node, the bundle passes into and of the muscular septum.
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*:N

- C.
I
- D.

-B.

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8It.. 0)utflos channiel ofI righIt seetricle. A. other e nd of t-Ot
(Fig. 7) seeni emerging betwseen thie septal cusps of' thle
ptIlranairtrsive. Thle dotted iennark s thc line of U'!lstIot
of the bulbar ridiges. and the site of type (ao dlet-ects as seen
trom right side. B. septal cusp oif the tricUspid valvse its
anterior edge hias beenl retracted to show the site otf the
interventricular part the patrs memntranacea septi. C
of

the cut Lirface otf the cristai suprasentricularis alt its

junction swith the ii erventlirtilar setptum. C). otiUS
rI SCle

Right Branch.-The right branch is a direct end. It usually forms a flat subendocardial band
continuation of the bundle and curves down- of parallel fibres of paler colour than the true
wards towards the apex, eit-her subendocardially myocardial tissue and running in a different direc-
or buried in the muscle. In this part of its course tion. The fibres fan out and two bands can some-
it runs dorsal to the conus muscle and in some times be seen, one going to the neighbourhood of
hearts can be traced into the moderator band. each papillary muscle.
Left Branch.-The left branch arises from the DEVELOPMENT
atrioventricular bundle at varying sites from the The conducting tissue can be recognized as
point where it emerges from the trigonum to its differentiating from the cardiac musculature
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310 E. W. T. MORRIS

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* ... ..^
., Or '.
Cv }w
kr Ei.\;

NNw f ,<d;;..
:S ,4
... Wi*; .:

FIG. 9.-10 mm. pig embryo. Transverse section. A.T., atria. V.V., ventricles. I.A.S., interatrial septum.
M.S., muscular part of the interventricular septum. X, region of the fused atrioventricular endocardial
cushion which forms the atrioventricular part of the pars membranacea septi.

during the sixth week (C.R.L. 7-9 mm.) and 'is complete and is then related to the posterior
rapidly extends, then the atrioventricular node is and inferior borders of the foramen which still
recognizable in the posterior wall of the atrium exists.
and the bundle extends along the dorsal wall of TYPES OF UNCOMPLICATED DEFECTS
the atrioventricular canal under the dorsal atrio-
ventricular cushion into the upper border of the These may occur in the following sites:
dorsal horn of the muscular septum, where it (a) In the septum separating the aortic and pul-
divides into a right and left branch. It is there- monary outflow, i.e., in that part of the septum
fore in situ before the final stage of the septum formed from the bulbar ridges; as seen from the
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THE INTERVENTRICULAR SEPTUM 311
right ventricle they occur above the crista supra- monly found are those with which a ventriculat
ventricularis and from both aspects they are situ- septal defect is combined to make up the clini-
ated.below the commissure of the septal cusps of cal entity of Fallot's tetralogy. In 120 cases of
the semilunar valves. Their site is therefore con- ventricular septal defect in the series of Warden
sistent with the bulbar ridges not fusing. et al., 45 were associated with other structural
abnormalities of the heart. In 33 (73.3%) of these
(b) In the region of the interventricular part of cases the associated defects completed the com-
the pars membranacea septi and adjacent muscu- plex of Fallot's tetralogy. In this condition the
lar septum: in general these defects as seen from division of the outflow channel has been com-
the right ventricle lie between the attachment of pleted by the fusion of the bulbar ridges. The
the septal cusp of the tricuspid valve and the septal stage of development of the interventricular sep-
attachment of the crista supraventricularis. tum which is at fault in this condition is the final
The crista supraventricularis therefore separates stage and the type of interventricular defect associ-
type (a) from type (b), the former lying above and ated with it is therefore class (b). It seems reason-
the latter below and behind it (Fig. 8). able to suggest that the final closure of the tem-
porary interventricular foramen by tissue derived
(c) In the muscular part of the septum: these from the dorsal atrioventricular cushion repre-
usually lie near the apex and are sometimes mul- sents a critical stage in the development of the
tiple, but may occur in the upper part of the heart. Its failure leads not only to the commonest
septum posteriorly, under the posterior part of type of uncomplicated interventricular septal de-
the septal cusp of the tricuspid valve. In this fect, but also contributes to the most commonly
purely muscular defect the atrioventricular bundle occurring complex of defects (Fallot's tetralogy).
and branches will probably lie anterior to and In this condition the other defects arise at a later
above the defect. stage in development than the septal defect which
ORIGIN OF THE DEFECTS.-By analogy with the accompanies them.

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known aetiology of congenital defects of other
parts of the body it seems likely that septal de- SUMMARY
fects may result either from failure of fusion of The steps in the development of the inter-
the several parts contributing to the septum, or ventricular septum have been described. Inter-
from a breakdown of the tissue once formed. It ventricular defects have been simply classified,
seems reasonable to assume that type (c) is formed and the relevant anatomical features of the
by the latter method. Of types (a) and (b), it is ventricles have been described.
impossible to say more than that they are in sites
compatible with a failure of fusion. The possible relationship between the types of
defect described and the steps in the development
FREQUENCY OF DIFFERENT TYPES OF DEFECT of the interventricular septum have been indicated.
In a series of 156 cases of ventricular septal Finally, other congenital defects of the heart
defects reported by Warden, DeWall, Cohen, frequently associated with interventricular septal
Varco, and Lillehei (1957) and Kirklin, Harsh- defects are briefly discussed. A short bibliography
barger, Donald, and Edwards (1957) six (3.8%) is given which should enable any aspect to be
were of type (a) and 145 (91.7%) were of type (b). followed up in more detail.
Class (b) is thus the most common defect in I wish to express my gratitude to Professor D. V.
these series, and this is what might be expected Davies for help and advice, and to Miss Dew for the
from a consideration of the stages in the develop- drawings in Figs. 1 to 6 and to Mr. A. H. Wooding
ment of the septum. This class of defect is com- for the photographs of Figs. 7 to 9.
patible with a failure of the final stage in the
formation of the septum, that is, the closure of BIBLIOGRAPHY
the temporary interventricular foramen by a EMBRYOLOGY
growth of tissue from the dorsal atrioventricular Arey, L. B. (1954). Developmental Anatomy, 6th ed. Saunders,
Philadelphia.
cushion, and it is in the final stages of a complex Frazer, J. E. (1953). Manual of Embryology, 3rd ed. Bailliere,
Tindall and Cox, London.
embryological process that developmental errors Hamilton, W. J., Boyd, J. D., and Mossman, H. W. (1952). Human
most commonly occur. This leads to a further Embryology, 2nd ed. Heffer, Cambridge.
Kramer, T. C. (1942). Amer. J. Anat., 71, 343.
reflection, namely, that when there are associated Odgers, P. N. B. (1938). J. Anat. (Lond.), 72, 247.
Tandler, J. (1912). Manual of Human Embryology, Vol. 2, ed.
congenital defects of the heart the ones most com- Keibel, F., and Mall, G. P. Lippincott, Philadelphia.
2C
 Thorax: first published as 10.1136/thx.12.4.304 on 1 December 1957. Downloaded from http://thorax.bmj.com/ on September 19, 2023 by guest. Protected by
312 E. W. T. MORRIS
ANATOMY SEPTAL DEFECTS
Johnston, T. B., and Whillis, J. (1954). Gray's Anatomy, 31st ed. Abbott, M. E. (1936). Atlas of Congeniital Car-diac Disease. Ameri-
Longmans, Green, London. can Heart Association, New York.
Walmsley, T. (1929). In Quain's Elemtzents of Anatomtiy, 11th ed., Becu, L. M., Fontana, R. S., DuShane, J. W., Kirklin, J. W., BurchelJ.
Vol. IV, Pt. Ill. Longmans, Green. London. H. B., and Edwards, J. E. (1956). Circulation, 14, 349.
Kirklin, J. W., Harshbarger, H. G., Donald, D. E., and Edwards,
CONDUCTING SYSTEM J. E. (1957). J. thorac. Surg., 33, 45.
Field. E. J. (1951). Brit. Heart J., 13, 129. Rokitansky, C. von (1875). Die Defecte der Scheidewainzde des
Herzens. Braumuller, Vienna.
Muil, A. R. (1954). J. Anat. (Lond.), 88, 381.
Walls, E. W. (1945). Ibid., 79, 45. Spitzer, A. (1923). Virchows Arch. path. Anat., 243, 81.
-- (1947). Ibid., 81, 93. Warden, H. E., DeWall, R. A., Cohen, M., Varco, R. L., and Lillehei,
Widran, J., and Ltv, M. (9151). Circdaltion, 4, 863. C. W. (1957). J. thoroc. Surg., 33, 21.

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