Anatomy and Pathology of the Whale Heart with

Special Reference to the Coronary Circulation

R. C. TRUEX, F. G . NOLAN, R. C. TRUEX, JR., H . P. SCHNEIDER
H. I. PERLMUTTER
Department o f Anatomy and Cardiovascular Research Institute of
Hahnemann Medical College, Philadelphia, Pennsyluania, and
The Marine Heart Research Foundation,
Los Angeles, California

The unique position of the whale in the

animal kingdom has long stimulated the
curiosity and challenged the research interest of the scientific world. These enigmatic mammals, whose remote ancestors
lived on land, have, during the evolutionary process of adapting to a marine environment, modified some of their organ
systems. It is not surprising, therefore,
that observations have been made upon
the anatomy, pathology and physiology of
the cetacean circulatory and respiratory
systems.
Knox (1838) observed the great size of
the heart in one adult and one fetus of the
fin whale (Balaena borealis Knox). He
stated the adult heart equalled in size an
ordinary sized washing-tub, while the
fetal heart was peculiar in form, being
much flattened, and scarcely exhibiting
the appearance of an apex. Pouchet and
Beauregard ( 1889, 1892) called attention
to the completion of an arterial circle on
the ventral surface of the cardiac infundibulum in two young whales (Balaenoptera
musculus and rostrata). These authors
also noted anastomoses between the marginal arteries and the ventral interventricular arteries in both specimens. White and
Kerr (15-27) examined the coronary vessels in the heart of an adult sperm whale,
but found no evidence of coronary artery
disease. Walmsley (38) exposed and measured the outside diameters of the coronary
arteries in a 320 pound heart of an adult
female fin whale (Balaenoptera physalus).
He also gave an excellent description of
the heart and coronary circulation in a fin
whale fetus with a body length of 143 cm.
Walmsley stated that the coronary veins
have the usual mammalian distribution

AND

although they are typical of the cetacean

venous system in that multiple channels
replace veins which are single in most
mammals.
Race, Edwards, Halden, Wilson and Luibe1 (59) recently completed a study on the
aorta, coronary arteries, venous drainage,
and cardiac muscle fibers in a 256-pound
heart of an adult male sperm whale
(Physeter catodon). They also studied the
heart of a sperm whale fetus with a body
length of 2.1 meters. These authors measured the adult heart and coronary circulation and graphically illustrated their results. Although they observed large, multiple anastomoses between the left and
right coronary arteries, they stated they
found no evidence of atherosclerosis or
calcification in any arteries.
The cardiac conduction system of the
whale has been examined morphologically
by several investigations (White and Kerr,
15-77; Ogata, 17; Ogata and Sassa, 19;
Walmsley, 38; Race et al., 59; Truex, 61).
Electrocardiograms have been recorded
from the live Beluga, fin, gray, and pilot
whales (King, Jenks and White, 53; White,
King and Jenks, 53; Kanwisher and Senft,
60; Nolan and Truex, unpublished data).
For physiological experiments on smaller
aquatic mammals the reader is referred to
the interesting papers of Irving (39),
Scholander (4O), Scholander and Irving
(41), and Irving, Scholander and Grinnell (41).
The above anatomical and pathologic
investigations on the whale heart and coronary circulation were of necessity based
Supported by U.S.P.H. grants H-1575, H-3816,
H-4753 and the Marine Heart Research Foundation.

325

326

R. C. TRUEX AND OTHERS

fusion method shown in figures 3 and 4.

Structures within the atria and ventricles
were exposed with little effort and are of
phenomenal size. However, this early experience with large hearts weighing up to
750 pounds convinced the authors it would
be prudent to concentrate their energies
on the smaller hearts of the sei and sperm
whales.
Caliper measurements of all the cardiac
valves and papillary muscles were completed, but for the sake of brevity, are
MATERIALS AND METHODS
omitted from the essential data shown in
The present investigation is based upon figure 1. The animals were all young
10 adult whale hearts of three different adults although neither their exact ages
species : (sei, or Balaenoptera borealis; nor body weights were determined. In a
gray, or Eschrichtius gibbosus; and sperm, general way the 4 chambered cetacean
or Physeter catodon); 7 specimens were heart resembled that of the land mammals.
from male and three from female animals. The left and right atria were capacious and
The aorta, pulmonary and caval vessels their auricles had thin distensible walls
were Uniformly measured and trimmed reenforced by numerous, thick, pectinate
prior to weighing each heart. Six hearts muscle strands. The capacity of the right
were dissected as fresh specimens, three ventricle approximates that of the left venafter formalin perfusion, and one heart tricle, while the most unexpected observawith latex injected into the coronary ar- tion was the variable thicknesses of the
teries. Variable amounts of fat ( 15 to 40 ventricular myocardium (fig. 1). In view
pounds) were carefully removed from the of the enormous work demanded of the
atrioventricular and interventricular sulci whale heart under several atmospheres of
to expose all of the epicardial branches of environmental pressure, we had anticithe two coronary arteries and the cardiac pated a uniformly thick ventricular wall.
veins. Each of the major arteries and veins Contrary to our expectations, the anterior
as well as their smaller rami were opened and lateral walls of the two ventricles as
and inspected for evidence of disease. The well as their apices, had areas that were
pattern of the coronary circulation of each surprisingly thin, whereas the myocardiumi
heart was diagrammed, after which the 4 was thick at the atrioventricular junction
chambers were opened, inspected and and within the muscular interventriculai:
measured. The essential measurements septum. Each whale heart, like their disand data for each of the 10 hearts are pre- tant ungulate cousins on land, had a heavy
muscular moderator band that spanned
sented in figure 1.
the lumen of the right ventricle (fig. 5).

upon a limited number of suitable specimens, often obtained by chance and studied
under difficult circumstances. Early in
1959 we had the good fortune to obtain
numerous fresh hearts from several species
of whales taken for scientific and commercial purposes. The hearts of several
fetuses, 6 adult fin, and three pilot whales
were partially dissected and studied, but
are not included in measurements reported
in the present communicatjon.

RESULTS

The cetacean heart is somewhat flattened when removed from the body and
emptied of blood, and is globular in shape.
The dorsal surface of the gray, sei, and
sperm whale hearts preisented a distinct
interventricular cleft. In addition the heart
of the sperm whale exhibited a bifid apex.
Preliminary studies on the very large hearts
of the gray and fin whale (figs. 2, 3, and 4)
were engrossing and informative, but physically exhausting due to the bulk of these
fresh specimens. The pattern and distribution of the coronary vessels were easily
determined by the crude but effective per-

Coronary circulation
The left coronary artery was dominant
in both gray whales and one sperm whale.
The right coronary artery was dominant
in one sperm whale heart and in another
was the only coronary artery arising from
the aorta. In the remaining 5 hearts the two
coronary arteries were of equal size and distribution. As in most mammalian heartis,
the left coronary artery of the whalle
provided nourishment to the left atriuin
and ventricle, while the right atrium arid
ventricle were supplied by the right coronary artery (figs. 6 and 7). Both coronairy

327

ANATOMY-PATHOLOGY O F WHALE HEART

ANATOMIC MEASUREMENTS OF 10 WHALE HEARTS
SPERM WHALE
(Physeter cotodon)
Body Length (in meters)------------13,1
Heort Weight (in Ibs.1--------------Circumference Heort tin em)--------137
Diometer Pulmonary Artery (in mml-- -I 87
Pulmonory Wall Thickness (in mm)-----I2
Diameter Aorta (in mm I- - I 90
Aortic Wall Thickness (in mm)---------18
Diameter Right Coronory Artery (in mml--30
Diometer Left Coronory Artery (in mm)----32
Left Ventricle Thickness (in mrn)
Lateral wall
43
Atrioventricular Junction--78
Apex
24

- --- - -

_-_------------- - ---- __________-__------__

Right Ventricle Thickness (in mm)
Anterior Wall-- -_------------ 30
Atrioventricular Junction---- - --- - 68
Apex-- - _ _ -_ __ _----- -----I 2

L e f t A-V Orifice (in mm)----------Right A-V Orifice (in mm)---------263

Moderator Bond (in mm)
Length
Width
Thickness
-

188

_ _ _ _ ______--- ----- 206

_ _ _ _ _ ___----- ------ 68
_ _ _ _ _ _ __ _ _ - - _ - - - -23
Fig. 1

lo*

@ __
a #~ 5 9
2 0_
1 _3 _
_
~

2.3

12.5
143
175
262

11.6
113
150
200

45
49

I I
210
18
33
35

190
14
44

10.5
I59
158
238
I3
200
17
35
22

I69
13
187

11.9

I54

132
200
13
150
20
28
30

162
206
26
175
20
33
36

I70
I75
17
200
18
32
38

I76
2 50
18
265
I8
34
44

'50
165
!37
12
I87
19
33
36

60
83
25

53
90
16

60
84
24

90
90
26

67
58
24

45
59
21

47
80
23

44
88
25

42
71
30

25
70
15
200
250

15
60
13
250
285

23
78
23
307
306

60
82
24
400
360

50
45
24
210
254

24
32
22
244
254

26
43
17
275
250

43
38
14
250
28 1

26
34
12

280

112
90
38

232
75
28

255
78
25

230
70
30

254
64
50

207
78
25

262
66
30

147
66
40

I54
60
30

I70

18

II

-- -

I90

-----

Size of cardiac structures and vessels.

arteries provided a rich arterial supply to

the interventricular septum by means of
the large ventral and dorsal interventricular arteries. The parent arteries then
anastomosed with each other. Multiple
large intercoronary anastomoses ( 5 to 15
mm in diameter) were demonstrated dorsally in the atrioventricular sulcus (fig. 7),
while smaller anastomotic vessels were
present on the ventral surface of the pulmonary artery, and along the ventral and
dorsal interventricular sulci.
Careful dissection of the arteries over
the base of the pulmonary artery and along
the interventricular sulci revealed arteriovenous communications 2-8 mm in diameter. Such channels were located entirely
within the epicardium and were observed
in all 10 hearts. The largest and most
conspicuous arteriovenous channels were
noted in the hearts of the sperm whale. In
the one heart with latex injected into the
coronary arteries under minimal pressure,
the latex was observed in all of the arteries but also in the large veins that accompanied the arteries within the inter-

ventricular sulci and proximal ventral wall

of the right ventricle (fig. 17).
Gray whale. The basic coronary arterial
pattern shown in figures 6 and 7 was found
in the two hearts of the gray whale. The
participating arteries were smooth walled
and devoid of any evidence of atherosclerosis or calcification. The ventral interventricular and right marginal arteries
(fig. 6 ) have counterparts in the human
heart, while the left marginal artery may
correspond to an enlargement of one of the
left ventricular branches observed in the
human heart. Additional ventricular arteries were present on both the ventral and
dorsal surfaces of the left and right ventricles (figs. 6 and 7).
The venous drainage of the ventricular
myocardium is accomplished by large epicardial veins that lie adjacent to, and parallel the course of, the major arteries. The
plicated walls of the larger veins were extremely thin. Numerous anastomotic veins
of variable size were present at the apex,
and on both the ventral and dorsal surfaces of the ventricles as shown in figures

328

R. C. TRUEX AND OTHERS

8 and 9. It was not uncommon to observe

large veins (8-10 mm in diameter) that
emerged abruptly from the deeper layers
of the myocardium to terminate in the
interventricular and marginal epicardial
veins. The basic venous pattern (figs. 8
and 9) was similar in both gray whale
hearts. As in higher land mammals, most
of the interventricular septum, left ventricle, and the dorsolateral wall of the right
ventricle were drained by tributaries of the
coronary sinus. The ostium of the coronary sinus was of considerable size (fig.
4 ) and was located in the dorsal wall of
the right atrium. The ventral and lateral
walls of the right ventricle were drained
by two to 4 veins. These vessels penetrated deeply into the epicardial fat of the
atrioventricular sulcus and terminated in
the ventral wall of the right atrium above
the annulus fibrosus. Small veins from the
lower portions of the right and left atria
terminated in the circumflex tributaries of
the coronary sinus. The roof and interatrial surfaces of the right atrium presented ostia of several minimal cardiac
veins similar to those observed in the heart
of dog and man (Truex and Schwartz, '51;
Truex and Angulo, '52).
Sei whale. The smaller heart of the sei
whale had a coronary vascular pattern and
distribution similar to that of the larger
gray whale (figs. 6, 7, 8, and 9). The
coronary arteries possessed numerous interarterial anastomoses between the two
parent vessels, had small arteriovenous
communications, but showed no evidence
of arterial disease. The wide venous channels were placed deep in the epicardium of
the interventricular and atrioventricular
sulci, whereas the ventral and dorsal surfaces of both ventricles had a veritable
network of small, anastomosing veins. The
marginal and circumflex veins were present, but in all three specimens the dorsal
interventricular vein was the largest tributary of the coronary sinus.
S p e r m whale. The coronary circulation
of these 5 hearts presented the basic arterial and venous patterns shown in figures
10, 11, 12, and 13. However, each specimen possessed its own anatomic peculiarities and vascular variations. The coronary
arteries and their myocardial branches
were conspicuous in this species due to

their large size and convoluted appearance. Only the larger arteries were included in figures 10 and 11. Unfortunately
the thick arterial network to the interventricular septum could not be shown
adequately in these two views. Abundant
intercoronary anastomoses between the
left and right coronary arteries were present over the base of the pulmonary artery,
in the interventricular and atrioventricular
sulci, and at the b s d apex. We were impressed particularly by the frequency and
large size of the anastomoses between
major branches of the left coronary artery
in all 4 specimens (figs. 10 and 16). We
have not observed such prominent intercommunications between branches of one
coronary artery in any of the other aquatic
or land mammals studied to date. The
female sperm whale heart with only a
right coronary artery arising from the
aorta was a most interesting specimen for
similar anomalies are observed only rarely
in man. In this whale heart the single
coronary artery gave rise to circumflex,
interventricular and marginal arteries
(figs. 14 and 15). These branches, though
all arising from one parent vessel, formed
a basic pattern of cardiac distribution similar to that found in the other sperm whale
hearts that possessed both a left and right
coronary artery. The arteries were explored thoroughly, but no evidence of
atherosclerosis or calcification was found
in this specimen. As demonstrated in figure 17 and noted above, the sperm whale
hearts all had numerous arteriovenous
anastomoses. Atheromatous plaques were
present to a varying degree in each of the
hearts of the 4 male sperm whales as described below.
The venous drainage of the sperm whale
heart is shown in figures 12 and 13. The
marginal veins were smaller, while the
interventricular and left circumflex veins
were of considerable size. The network of
small epicardial veins observed in the
hearts of the sei whale was here replaced
by a more open network composed of larger
veins. This observation was in marked
contrast to our findings in the gray and sei
whales. The venous radicals in the heart
of the sperm whale were neither as large
nor as numerous as the arteries that supplied the myocardium.

ANATOMY-PATHOLOGY O F WHALE HEART

Pathology
The epicardial arteries of each heart
were opened and examined for evidence of
coronary artery disease. The intima and
walls of all of the arteries of the gray and
sei whales were smooth and devoid of disease. The aorta, coronary orifices, and
lumens of the major proximal branches of
the left and right coronary arteries of all
of the sperm whale hearts also appeared
smooth and entirely normal.
However, in the more distal portions of
the marginal, interventricular, and ventricular arteries of each of the 4 male
sperm hearts we observed multiple atheromatous intimal plaques. In one specimen
the plaques were of small size (fig. 19)
and located most commonly a,t the ostium
of a smaller branch of the parent vessel.
In two specimens multiple medium-sized
plaques were observed (figs. 21 and 22).
The elongated, yellowish, intimal plaques
were always oriented parallel to the long
axis of the artery. In no artery did we
observe an annular type lesion that formed
a ring within the lumen of the vessel.
Microscopic sections of the atheromatous plaques, prepared by a variety of differential stains, demonstrated a histological appearance (figs. 23 and 24) quite
similar to that observed in human coronary arteries with atherosclerosis. The
plaques were composed of proliferated tissues that involved chiefly the intima and
subintimal layers. The fibrous elements
predominated, particularly collagen fibers.
Numerous fibroblasts and macropha.ges
were distributed between the abundant
connective tissue fibers. Hemorrhage and
blood pigments often were observed within the confines of the larger plaques, although we observed no evidence of necrosis or calcification within the walls of the
arteries. The internal elastic membrane
had lost its integrity and was replaced by
a loosely fenestrated fibrous layer (fig. 23).
In some instances the inner layers of the
tunica media were also involved, and in
such regions there was no sharp demarcation between the tunica intima and tunica
media. The clear areas between the wavy
fibrous layers within the plaque, and along
the intimo-medial junction (fig. 24), resulted from the extraction of lipids during

3219

tissue dehydration and embedding. Frozan

sections stained with Scharlach R demomstrated an abundance of minute fat droplets within the plaques that were arranged
in stratified layers between the fibrous elements. Larger deposits of fat were also
observed within the plaques as well as
along the intimo-medial junction.
In only one specimen did we observe
evidence of myocardial infarction and necrosis. The heart of one sperm whale possessed a circumscribed lesion that had destroyed the proximal and dorsal portion of
the muscular interventricular septum and
adjacent wall of the left ventricle. Thk
entire thickness of the septum had been
replaced in an area of 10 X 15 cm by a
foul smelling mass of cheesy material interlaced by fibrous strands. The margins
of the necrotic area were not sharply demarcated from the cardiac muscle fibers
of the surrounding myocardium. The gross
pathologic appearance of the area was that
of a recent dorsal myocardial infarction
undergoing necrosis. In this specimen the
coronary arteries had demonstrated multiple medium-sized atheromatous plaques,
and also nematode worms had been found
in the cardiac veins (figs. 5 and 18). We
were unable to isolate and identify the
specific artery, or arteries and veins, responsible for the infarcted area in this
heart.

Parasitism
Examination of the small proximal veins
on the dorsal surface of two sperm whale
hearts revealed the presence of nematode
worms. The lumen of one vein was entirely occluded by one large worm that was
2.5 mm in diameter and 125 mm in length
(fig. 18). The right ventricle of this same
specimen contained 15 additional unidentified nematode worms, 90 to 129 mm in
length (fig. 5).
DISCUSSION

Although gross body measurements and

organ weights have been made in several
species of whales, little scientific information has accumulated concerning the heart
and circulation of these mammals. To the
authors knowledge this is the only study
based upon several whale hearts. We have
recorded some of the cardiac measure-

330

R. C. TRUEX AND OTIIERS

ments (fig. 1 ) and presented the basic

coronary vascular patterns (figs. 6 to 13)
of the whale as only a starting point for
future investigations on additional animals.
The homologies between the coronary
circulation of the cetacean and human
hearts can be appreciated best by a careful examination of these illustrations. For
example, the ventral interventricular and
left circumflex arteries have a n origin and
distribution similar to the anterior descending and circumflex branches 0f the human
left coronary artery. In a similar manner
the left circumflex vein corresponds to the
great cardiac vein of the human heart; the
dorsal interventricular vein corresponds to
the middle cardiac vein, while the right
marginal vein is analogous to the small
cardiac vein of man. The right ventricular
veins, though few in number, simulate the
human anterior cardiac veins, but in the
cetacean heart they possess more extensive anastomoses with the large tributaries
of the coronary sinus.
The greatest difference between the
whale and human coronary circulation is
manifest in the multiple, large intercoronary arterial anastomoses, anastomoses between large branches of one coronary artery, and the unusual venous drainage of
the whale heart. One can demonstrate
small intercoronary anastomoses on the
dorsum of the human heart, but never
finger-sized vessels comparable to those
always observed in the same region of the
whale heart. One can only speculate
whether such interarteria'l anastomoses
are of small caliber in man because of the
smaller size of the human heart and coronary arteries as compared to these structures in the whale. The large anastomoses
between the tortuous major branches of
the left coronary artery of the sperm
whale are unique even among the whales,
and are normally absent in the human
heart.
The occurrence of arteriovenous anastomoses in all of the whale hearts merits
additional consideration. The size, location and frequency of such anatomic communications indicates they must have
some important, but still unknown, functional significance. One can only surmise
that they may play an important role in

the shunting of arterial blood into the

venous system during myocardial engorgement or distention. It is well known that
both the baleen and toothed whales exhale carbon dioxide and take in atmospheric air when they surface to "b10~"
several times. Blow holes, located on the
top of the head, remain tightly closed
when the whale is submerged, and the
animal must perform all of his bodily
functions on stored oxygen. It is possible
that venous blood may cause distention
and engorgement of the right ventricle
and right atrium during prolonged submersion. Under such circumstances the
ventricular walls would be compressed,
and would offer increased resistance
to coronary arterial flow. Under these
circumstances the arteriovenous shunts
might act as safety valves that direct the
arterial blood into the low pressured venous system.
Comparative aspects of the coronary
circulation merit a more extensive study,
for several authors have recently observed
anomalous arteriovenous communications
in the human heart (Walther, Starkey,
Zervopolus and Gibbons, '57; Schultz, '58;
Steinberg, Baldwin and Dotter, '58; Edwards, '58; Zuhdi, Kraft, Carey and Allen,
'60). Such anomalous human arteriovenous anastomoses correspond in location to
those we have observed in the normal coronary circulation of the whale heart. The
phylogenetic significance and possible relationship between the arteriovenous anastomoses observed in the aquatic and land
mammals including man, must await further investigation.
Marine nematodes have been found in
the hearts of several aquatic mammals.
Heart and lung worms most commonly
are observed in those aquatic mammals
that live on a fish diet such as seals, sea
lions, and dolphins. (Fasut, '37; Lyster,
'40; Dougherty and Herman, '47; Anderson, '52; Margolis, '54; Brown, McIntyre,
DelliQuadri and Schroeder, '60). Nematodes have been recorded previously in
the right ventricle of the sperm whale
heart (see Margolis, '54), but to our knowledge they have never before been observed
within the epicardial cardiac veins. Parasitic nematodes in the right heart could
migrate with relative ease into the ostium

ANATOMY-PATHOLOGY OF WHALE HEART

of the fist-sized coronary sinus and its

large venous tributaries during atrial diastole.
We believe the existence of atheromatous plaques in the coronary arteries of the
male sperm whale to be a most significant
finding. Future studies, on a more extensive series of whale specimens of both
sexes, must be made before one can determine the validity of such factors as
age and sex upon atherosclerosis. However, it should be noted that the sperm
whale dives deeper, remains submerged
longer (50-60 minutes), has a squid and
fish diet, and an erratic temperament that
set him apart from the other species of
whale. One can only speculate as to the
special roles played by stress, diet, and
temperament in these toothed denizens of
the deep. We look forward to the examination of the coronary arteries of truly
aged sperm whale bulls. Such aged bulls
live an isolated life in the frigid arctic
and antarctic waters, and presumably
have been driven from the herds by
younger males. Time and study alone will
reveal whether such aged animals possess a more advanced degree of coronary
atherosclerosis.
We cannot explain the absence of
plaques in the coronary arteries of the
sperm whale as reported by White and
Kerr (15-17) and Race et al. (59). The
presence of atherosclerosis in our 4 male
specimens may have resulted from a fortuitous sampling of specimens, or a more
thorough examination of all the distal arterial branches. We reiterate, the intimal
atheromatous changes were always confined to those distal arteries that feed
branches directly into the ventricular myocardium. Unless these small arteries are
opened and examined with care one
would miss the vascular pathology described and illustrated in this communication. An extensive search will be made
for evidence of myocardial infarction in
our continuing studies.
We realize we have created as many
puzzling problems as we have solved.
We have derived some solace from the
knowledge that many investigators who
studied whales before us have often been
amazed, bewildered, and perplexed by
seemingly unsolvable questions. We sub-

331

scribe whole-heartedly to the words of a

noted predecessor in this field, P. D. Scholander (59) who stated: When playing
around in the ocean, dolphins are pleasing to the eye no end, but let it only add
to your thrill that these rascals are a
graveyard to our wits. For is not finding
out infinitely more exciting than knowing
the answer?
SUMMARY AND CONCLUSIONS

The present study of the adult whale

heart and coronary circulation was made
upon 10 specimens of three different species (sei, gray, and sperm whales) with
heart weights that ranged from 100 to
350 pounds. Branches of the coronary
arteries and cardiac veins were dissected,
diagrammed and examined for evidence
of arterial disease. Essential measurements of the heart and blood vessels are
presented, while the basic patterns of the
cardiac arterial and venous systems are
illustrated and compared to those of man.
The right and left coronary arteries
were of equal size and distribution in 5
hearts; the left coronary was dominant in
three, and the right coronary artery was
dominant in one specimen. An anomalous
right coronary artery was the only coronary vessel present in one sperm whale
heart. Large intercoronary anastomoses
between the left and right coronary arteries were demonstrated in 9 hearts,
while anastomoses between the major
branches of the left coronary artery were
a prominent feature of the circulation in
the heart of the sperm whale. Arteriovenous communications were observed in
all 10 specimens, and provide a possible
phylogenetic explanation for the troublesome arteriovenous anomalies recently encountered in the human coronary circulation. Marine nematode heart worms were
observed in the right ventricle and cardiac
veins in two of the sperm whale hearts.
One male sperm whale heart demonstrated a large recent myocardial infarct
in the interventricular septum. The significance of these results are discussed.
The coronary arteries of the sei and gray
whales were devoid of atherosclerosis and
calcification. However, we observed multiple, yellowish colored, intimal atherom
atous plaques in the more distal ventric-

332

R.

C . TRUEX

ular branches of both coronary arteries in

the hearts of the 4 male sperm whales.
The factors of sex, age, species, stress, and
diet appear to be strongly implicated in
cetacean arterial atherosclerosis, but the
significance of each factor must await
more adequate sampling and future investigation.

AND OTHERS

Irving,,L., P. F. Scholander and S. W. Grinnell

1941 The respiration of the porpoise, Turaiops truncatus. J. Cell. and Comp. Physiol.,
17: 145-168.
Kanwisher, J., and A. Senft 1960 Physiological measurements on a live whale. Science,
121: 1379-1380.
King, R. L., J. L. Jenks and P. D. White 1953
The electrocardiogram of a Beluga whale. Circulation, 8: 387-393.
Knox, F. J. 1838 Catalogue of Anatomical
ACKNOWLEDGMENTS
preparations illustrative of the whale. Particularly the Great Northern Rorqual (Balaencx
The authors are indebted to the U. S .
Maximus borealis). Neill and Co., Edinburgh,
Department of Interior, Fish and Wildlife
pp. 1-38.
Service for special permission to euthanize Lyst&r,L. L. 1940 Parasites of some Canadian
sea mammals. Canad. J. Res., 18: 395-409.
and study the California gray whale; to
Charles and John Caito of the Del Monte Margolis, L. 1954 List of the parasites recorded from sea mammals caught off the west
Whaling Station, Richmond, California
coast of North America. J. Fish. Res. Bd. Canfor laboratory facilities and technical asada, (Pacific Biolog. Sta., Nanaimo, B.C.),
sistance; to the British Columbia Packers,
1 1 : 267-283,
Whaling Division, Vancouver and Coal Ogata, T. 1917 Uber die Struktur des Walfischherzens, besonders seines Reizleitungssystems
Harbour, B. C., for their valued assistance
(I. Mitteilung). Jap. Path. Gesellsch., 7: 87-88.
in providing laboratory space and fresh
Ogata, T., and K. Sassa 1919 Uber das spezispecimens for this study; to Carlernst
fische Reizleitungssystem im Vorhof des WalDiedrich, Bruce McNeil, Lindsley Parsons,
fischherzens. Jap. Path. Gesellsch., 9: 3637.
Hector Cowie, Hanibul Anderson, Captain Pouchet, G., and H. Beauregard 1889 Recherches sur le cachalot. Nouv. Arch. Mus.
Edward Karlsen, Captain Arne Borgen,
Hist. Nat. Paris, ser. 3, vol. I, pp. 1-96.
Richard Cohen, Faith Hartman, Martha Q.
1892 Recherches sur le cachalot (suite).
Smythe, Audrey Gourley, Theresa MalinIbid., vol. 4,pp. 1-90.
owski, and Ilona Bauer for their coopera- Race, G. J., W. L. J. Edwards, E. R. Halden,
tion and professional assistance.
H. E. Wilson and F. J. Luibel 1959 A large
whale heart. Circulation, 19: 928-932.
We express our sincere appreciation to
Miss Marjorie Stodgell, Steven Gigliotto, Scholander, P. F. 1940 On the respiratory adjustment to prolonged diving in the seal. Am.
Louis Sunny, and Walter Guzejko for their
J. Physiol., 129: 456457.
valued medical art and photographic as1959 Wave riding dolphins, by W. D.
sistance in preparing the illustrations used
Hayes. A reply. Science, 130: 1638.
in this publication.
Scholander, P. F., and L. Irving 1941 Experimental investigations on the respiration and
diving of the Florida Manatee. J. Cell. and
LITERATURE CITED
Comp. Physiol., 17: 169-191.
Anderson, R. C. 1952 Description and relationSteinberg, I., J. Baldwin and C. Dotter 1958
ships of D i r o f i h i a Ursi Yamaguti 1941, and a
Coronary arteriovenous fistula. Circulation,
review of the genus Dirofilaxia. Trans. Roy.
17: 372-390.
Canad. Inst., 29: 35-65.
Truex, R. C. 1961 Comparative anatomy and
Brown, D. H., R. W. McIntyre, C. A. DelliQuardi
functional considerations of the cardiac conand R. J. Schroeder 1960 Health problems
duction system. Rio de Janeiro Symposium:
of captive dolphins and seals. J. Am. Vet.
Electrophysiology of specialized cardiac fibers.
Med. Assoc., 137: 534-538.
Elsevier Publ. Co., Amsterdam, (in press).
Dougherty, ,E. C., and C. M. Herman 1947 Truex, R. C., and A. W. Angulo 1952 ComNew species of the Genus Parafilaroides,
parative study of the arterial and venous sysDounhertv. 1946 (Nematoda: Metastronnvlitems of the ventricular myocardium with spedae< Pinkpedia. 'Proc. Helminth. SOC. of
cial reference to the coronary sinus. Anat.
Wash., 14: 77-87.
Rec., 113: 467-492.
Edwards, J. E. 1958 Anomalous coronary arTruex, R. C., and M. J. Schwartz 1951 Venous
teries with special reference to arteriovenous
system of the myocardium with special refercommunications. Circulation, 17: 1001-1006.
ence to the conduction system. Circulation,
Faust, E. C. 1937 Mammalian heart worms of
4: 881-889.
the genus Dirofilaria. Festschrift Bernhard
Walmsley, R. 1938 Some observations on the
Nocht., 80: 131-139.
vascular system of a female fetal finback.
Irving, L. 1939 Respiration in diving mamContr. Embryol. Carneg. Instn., 37: 109-192.
mals. Physiol. Rev., 19: 112-134.

-_

ANATOMY-PATHOLOGY O F WHALE HEART

Walther, R. J., G. W. Starkey, E. Zervopolus and

G. A. Gibbons 1957 Coronary arteriovenous
fistula: Clinical and uhvsiologic reuort of two
patients, with review of the- literature. Am.
J. Med., 22: 213-222.
White, P. D., and W. 3. K e n 1915-1917 The
heart of the suerm whale with esuecial reference to the A-V conduction system. Heart,
6: 207-216.

333

White, P. D., R. L. King and J. L. Jenks 1953

Thesrelation of heart size to the time intervals
of the heart beat. with uarticular reference
to the elephant and the whale. New Eng. J.
Med., 248: 69-70.
Zuhdi, N., D. Kraft, J. Carey and A. Greer 1960
Coronary arteriovenous-like communications.
A.M.A. Arch. Surg., 80: 178-180.

PLATE 1
EXPLANATION OF FIGURES

334

Ventral surface: 250-pound heart of a male gray whale. A hypertrophied human heart
of 400 gm is superimposed on the right atrium to illustrate comparative heart slze
of the two specimens. The major arteries and veins are concealed by large amounts
of fat in the atrioventxicular and interventricular sulci as well as along the margins of
both ventricles. The letters used in all figures indicate the 4 heart chambers (Le.,
RA, right atrium; RV, right ventricle; LA, left atrium; LV, left ventricle).

Distention of myocardium of right ventricle by two inch hose inserted i n right coronary artery of fresh fin whale heart (300 lbs.). The left ventricle could be distended
i n a similar manner .with water pressure from a three inch hose placed in the left
coronary artery. This effective method was used to demonstrate both the arterial and
venous patterns of several fresh specimens.

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 1

335

PLATE 2
EXPLANATION OF FIGURES

336

Ostium of coronary sinus ( C S ) in the right atrium of a fin whale heart weighing
300 pounds. Openings of the three largest venous tributaries ( f i g s . 9 and 13) of the
coronary sinus are clearly shown. Venous return is illustrated during irrigation of
myocardium with equal pressures in the left and right coronary arteries. The right
atrioventricular orifice and posterior papillary muscle of the right ventricle are seen
on the Tight side of the figure.

Right ventricle of sperm whale heart (specimen number two, i n fig. 1). The coronary
circulation of this specimen is shown i n figures 10, 11, 12, and 13. Marine nematodes
were present i n the right ventricle ( F and G , fig. 5 ) , and cardiac veins (fig. IS) of this
heart. The specimen also demonstrated a recent, large infarction of the interventricular
septum. The anterior ( A ) and posterior ( C ) papillary muscles; moderator band ( B ) ;
papillary muscle of the conus (D); and valves of the pulmonary artery ( E ) are identified.

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G . Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 2

337

PLATE 3
EXPLANATION OF FIGURES

338

Coronary arteries on ventral surface of a 250-pound gray whale heart. The major
arteries are identified and their respective diameters are indicated.

Coronary arteries on dorsal surface of gray whale heart. Same specimen as shown in
figure 6 above. Note intercoronary anastomoses in dorsal atrioventricular and interventricular sulci, and a t apex.

PLATE 3

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G . Nolan. R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

R. Ventricular
(7-13 rnm)

R. Marginal
(12 mm)

Ventral Interientricular a.
(20-28 mm)

339

PLATE 4
EXPLANATION O F FIGURES

8 Cardiac veins on ventral surface of a 250-pound gray whale heart. The major veins
are identified and their respective diameters are indicated. Anastomoses between the
ventricular veins and coronary arteries (fig. 17) were present i n all the hearts of the
gray, sei, and sperm whales.

9 Cardiac veins on the dorsal surface of gray whale heart. Same specimen as shown in
figures 6, 7, and 8. Note large size of dorsal interventricular vein and ostium of
coronary sinus.

340

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G . Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 4

Aorta

Circumflex
(24 mm)

L. Marginal v.
(16-20 mrn)

Dorsal 1nt;rventricular
(27-40 mm)

v.

9
341

PLATE 5
EXPLANATION O F FIGURES

10 Coronary arteries on ventral surface of a sperm whale heart. The tortuous major
arteries are identified, and the prominent intercoronary anastomoses are clearly shown.
The sperm whale hearts also possessed large anastomotic communications between
major branches of the left coronary artery (fig. 16).

11

342

Coronary arteries on dorsal surface of a sperm whale heart. Same specimen as shown
in figures 5 , 10, 12, arid 13. This heart also demonstrated a recent large infarct in
the dorsaI and proximal part of the interventricular septum. Note bifid apex of the
sperm whale heart.

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 5

a.

cular aa.

I0

343

PLATE 6
EXPLANATION OF FIGURES

12 Cardiac veins on ventral surface of sperm whale heart. The major veins are identified.
The right ventricular veins terminate in the right atrial cavity and also have large
communications with the tributaries of the coronary sinus.
13 Cardiac veins terminating i n coronary sinus on dorsal surface of sperm whale heart.
Although there are variations i n vessel size and their arrangement, the basic pattern
of venous drainage is similar i n the three species of whales studied (e.g., compare
figs. 8 and 9 with figs. 12 and 13).

344

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 6

nflex v.

vv.

12

345

PLATE 7
EXPLANATION O F FIGURES

14 Anomalous coronary circulation of heart. The heart of this female sperm whale had a
single right coronary artery that supplied the myocardium of all 4 chambers. Although
arising from a single coronary artery, the major branches conform to the basic arterial
pattern and distribution observed in other whale hearts (e.g., compare figs. 14 and 15
with the arterial patterns shown in figs. 6, 7, 10, and 11).
15 Branches of a single right coronary artery on the dorsal surface of sperm whale heart.
Same specimen as shown i n figure 14.

346

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 7

a.

Circumflex a

Marginal a.

15
347

PLATE 8
EXPLANATION OF FIGURES

16

Anastomoses between major branches of the left coronary artery of a sperm whale
heart injected with latex. The large latex communications between the ventral interventricular ( l ) ,ventricular, and circumflex ( 2 ) arteries are clearly shown after careful removal of the vessel walls. Large amounts of the thick latex, injected into the
arteries, were recovered in the adjacent more flattened cardiac veins (3).

17

Arteriovenous anastomoses on ventral surface of right ventricle of a sperm whale heart

with latex injection of coronary arteries. The auricle of the right atrium (1) and right
coronary artery ( 2 ) .are identified adjacent to the pulmonary artery. The smaller
arteries shown in the illustration communicated directly with the injected vein held in
the center of the figure. Such arteriovenous anastomoses were demonstrated in all
10 whale hearts, and were most often located over the base of the pulmonary artery,
and along the atrioventricular and interventricular sulci. Similar anastomoses, requiring surgical correction have been observed recently i n the human heart (see text).

18 Marine nematode obstructing the lumen of a cardiac vein (1) on the dorsal surface
of the left ventricle of a sperm whale heart. Same heart illustrated in figures 5, 10,
11, 12, and 13. The dorsal interventricular artery ( 2 ) is identified. Deep to the area
shown in this figure we observed a large, recently infarcted region within the interventricular septum.

348

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 8

349

PLATE 9
EXPLANATION O F FIGURES

19 Two small intimal atheromatous plaques (arrows) in a branch of the right marginal
artery of a male sperm whale. Multiple plaques were observed in many of the smaller
arteries over both ventricles i n the hearts of the 4 male sperm whale specimens.
Magnification of gross specimens in figures 19-22 is indicated by metric scale.

350

20

Medium-sized intimal atheromatous plaque (arrow) in a left ventricular artery of a

male sperm whale. Note proximity of plaque to the orifice of a smaller branch of the
parent vessel.

21

Large intimal plaque (arrow) in ventral interventricular artery of a male sperm whale.
The large plaques were always parallel to the long axis of the artery. No annular
intimal lesions were observed i n this study. Note abundant amount of periarterial
adipose tissue so common to the cetacean coronary circulation.

22

Multiple, large, intimal atheromata (arrows) in a branch of the dorsal interventricular

artery of a sperm whale heart. The lumen of the artery was partially occluded i n each
instance, however, complete occlusion of a n artery was not observed.

ANATOMY-PATHOLOGY OF WHALE HEART

R. C. Truex, F. G. Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 9

351

PLATE 10
E X P L A N A T I O N O F FIGURES

352

23

Microscopic section of intimal atheromatous plaque. Fibroblasts, macrophages, and

blood pigments account for the cellular and mottled appearance. Note absence of
internal elastic membrane. Male sperm whale. Hematoxylin and eosin stain. x 30.

24

Photomicrograph demonstrating fibrous structure of the intimal atheromatous plaque.

The darkly stained collagen fibers predominate within the plaque, tunica intima, deeper
layers of the tunica media, and tunica adventitia. Male sperm whale heart. Van
Gieson stain. >( 20.

ANATOMY-PATHOLOGY OF WHALE HEART

R. c. Truex, F. G.Nolan, R. C. Truex, Jr., H. P. Schneider and H. I. Perlmutter

PLATE 10

353