SOME OBSERVATIONS ON THE CETACEAN STOMACHS,
WITH SPECIAL CONSIDERATIONS ON THE
FEEDING HABITS OF WHALESHIROSHI HOSOKAWA* AND TOSHIRO KAMIYA**ABSTRACT

Macro- and microscopical observations on the sei, fin, blue and sperm
whale stomachs are described and discussed on the base of comparative-anatomical and functional viewpoint. The compartmentalization of the cetacean
stomach is a unique character in the animal kingdom and it must be attributed
to the aquatic life of the group. An attempt has been made to establish therelationship between the anatomical features of the cetacean stomach andits physiological function.
INTRODUCTION
In the latter half of the seventeenth century Tyson (1680) described the complicatedstomach of the common porpoise. About one hundred years later Hunter (1787) remarked that the cetacean stomach revealed generally a highly characteristic feature ofthis group of animals, divided into several compartments. He noticed, furthermore,that it resembled very closely those of cattles and sheep, and he supposed the whales
probably belonged to the ruminants, or had at least an intimate phylogenetical
relationship with the ungulata. Thereafter, many anatomists directed their attention to the cetacean stomach and numerous reports were published, dealing directly
or incidentally with this special feature of cetacean anatomy. Though it was soon
ascertained that the cetacean stomach was, in spite of its resemblance to the ruminantone, quite different from the latter, its extraordinarily complicated structure remained for a long time without a clear comparative-anatomical explanations, not tospeak of its physiological interpretation. The specifical differences in form of
stomachs among all different kinds of whales were so extreme that it was very
difficult to abstract the standard figure of them. We will quote here, for example,
various numbers of stomach-chambers recorded by many authors for Mystacoceti. In
most cases, the research materials of these authors were adult or fetal minke whale(Balaenoptera acutorostrata ), though some of them studied the blue whale (B. musculus)
or fin whale (B. physalus). Schulte alone used a fetus of the sei whale (B. borealis).
Hunter (1787) 5,

Cuvier (1805) 4,

Meckel (1829) 3,* The late Professor of the Department of Anatomy, Faculty of Medicine, University of Tokyo, he deceasedon the way to this work in 1967.

** Department of Anatomy, Faculty of Medicine, University of Tokyo, Tokyo.
Sci. Rep. Whales Res. Inst.,
No. 23, 1971. 91-101.

92

HOSOKAWA AND KAMIYA

Eschricht (1849) 3, Murie (1865) 4, Carte & Macalister (1868) 5,
Perrin (1870) 4, Weber (1888) 3*, Pilliet (1891) 4,
Turner (1892) 5, Schulte (1916) 4*, Slijper (1962) 3*.
*without counting the duodenal ampulla.
At the end of the nineteenth century, however, ·weber (1888) and Jungklaus(1898) published very splendid works on this problem and the macroscopic anatomyof the cetacean stomach was brought to a general conclusion. Thus, Weber classified the stomach of toothed whales, based on the development of its first compartment covered with the oesophageal mucous membrane, into the following 5 types:
3) Phocaena-type,1) Ziphiidaes-type,2) Hypothetic intermediate-type,4) Globicephalus-type and 5) Lagenorhynchus-type, and he explained thatthese types had developed one by one in the order of their number from the mostprimitive Ziphiidaes type. On the other hand, according to J ungklaus, who
endeavoured to use generally the comparative-anatomical characteristics of thecetacean stomach, not only of the toothed whales but also of the baleen whales, themost remarkable peculiarity of the cetacean stomach distinguishing itself from thesame of other mammals did not lie in its mere complexity caused by the compartmentalization but consisted in the fact that the glandular stomach itself was separated
into two divisions, one of those being provided with the peptic or fundus glands
and the other with the pyloric glands. The latter divisions were, moreover, apt
to be divided into two or more secondary compartments. On the difference between
the Odontoceti and Mystacoceti stomachs he stated that in the latter the constrictions
between each divisions or compartments were not so sharp as in the former. Jungklaus's conclusion of the characteristic feature for cetacean stomach is quite rightfrom the comparative-anatomical standpoint, as was confirmed afterwards by
Beddard (1900). From the physiological standpoint, however, we suppose that thecomplexity of the cetacean stomach, which is caused not only by the compartmentalization of the glandular portions but also by the oesophageal dilatation etc. has agreat significance. Against this background information we will, in the present
paper, try to present a detailed histological picture of the cetacean stomach.

MATERIALS

Baleen whale:
14.0 m male, 14.6 m female, 15.7 m female,
16.2 m female.
Fin whale (Balaenoptera physalus): 21. 3 m female.
Blue whale (Balaenoptera musculus): 23.8 m male.
Sei whale (Balaenoptera borealis):

Toothed whale:
Sperm whale (Physeter catodon): several adults, of which one 11.0 m longfemale and one 15.6 m long male were studied with special attention.
Sci. Rep. Whales Res. Inst.,
No. 23, 1971.

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94

HOSOKAWA AND KAMIYA

pancreas. The other two, namely the second and the third, are real gastric compartments providing with digestive glands. Between these two, however, there isanother small division developed especially in the fetal stage. In the female sei
whale of 15. 7 m length the largest compartment is the third one, measuring 110 cm
and 85 cm respectively in the transverse and longitudinal diameters in relation to
the axis of the digestive canal, followed by the fourth (85 x 75 cm) and the second(80 x 80 cm) in that order, and the first compartment is still smaller. Communication between the first and the second chambers is through a not so strong constriction. Where the second chamber leads into the third through the small connecting
division, there exist two semilunar valves separating the two compartments. The
third compartment opens into the fourth through the narrow passage constricted
sharply by the pyloric sphincter (Fig. 1).
Macroscopical features of the mucous membrane: (Plate I)
The mucous membrane of the oesophagus is lined by the thick, graywhite andcornified epithelium folded with numerous longitudinal, fine furrows. The sameepithelium clothes the inner surface of the first compartment too, and there aremany reticular folds of several millimeter in width, which are ramified and anastomosed to each other. These reticular folds disappear, when the wall of the stomach is
stretched, so that they are formed by the undulation of the whole wall. It is clear,
therefore, that this compartment is able to contract and dilate to a considerable
extent by the aid of the mural muscle strata. Where the first compartment leads
into the second one, the mucous membrane changes its character abruptly, there is adistinct lineal boundary between the oesophageal epithelium of the former and thegastric lining of the latter (Plate I-1 ). This is covered by the soft, mucous epithelium of dark gray colour with some tone of yellow. Its surface is smooth for 2-3 cmfrom the line separating the oesophageal epthelium and the gastric lining referred
to above, followed by thick reticular folds, about 1 cm in width, resembling theconvolution of the human cerebrum. These folds do not disappear even when thestomach is stretched; that is to say that they are formed in the mucous membrane
itself merely for the purpose to increase the inner surface of the stomach. The
second compartment has, therefore, no anatomical feature to contract or dilate
itself extensively. In the lower half of the second compartment the reticular folds
are reduced as the compartment approaches the third one, in which the surface ofthe mucous membrane is quite smooth and without any furrows or folds. It iscoloured yellowish gray, with some tone of pink. The duodenal ampulla is clothed
also with smooth, soft membrane, coloured brownish red and it continues with thatof the small intestine, without any constriction or valve. For the female sei whale(15.7 m long) the first compartment has the thickest wall measuring 2.5 (-3.0) cm,
containing very well-developed muscular strata. The wall of the remaining three
are somewhat thinner and measure about 1.2 (1.0-1.5) cm. The wall of the small
intestine is 0.8-1.0 cm thick. For the male blue whale of 23.8 m length the wallsof the first, second, third and fourth compartments were 4.0 cm, 2.0 cm, 1.5-2.0 cm(mucosa: 0.5, muscularis: 0.9, serosa: 0.1) and 1.3 cm (mucosa: 0.3, muscularis:
0.7, serosa: 0.3) respectively.
Sci. Rep. Whales Res. Inst.,
No. 23, 1971.

CETACEAN STOMACH AND FEEDING HABITS

95

Histological structure: (Plate II)
Oesophagus is lined by an extensively stratified flat epithelium of about 1 mm
in thickness. The most superficial layer of the epithelium (25-30 µ) is completely
cornified and the cells have no trace of nucleis in contrast to the case in humans.
The next layer of about 25 µ has cells containing rudimentary nuclei. These twolayers are followed by a third layer, 4-5 cells in thickness the cytoplasm of the cells
staining deep red with eosin and the nuclei are shriveled, indicating probably thecommencement of the cornification of cells. The next layer is composed of several
strata oflarge cells, occupying the greater part of the epithelium. While the cells of
the upper portion of this layer are situated with their axis almost parallel to thesurface of the epithelium, those of lower part make a right angle with the same especially in the areas between the papillae. While the nuclei of the former are
smaller and stain deeply, those of the latter are larger and stain lightly. The deepest
layer is bounding to the propria, and composed of crowded cells of small size. The
connective tissue, propria, of about 3-4 mm in thickness, contains many vessels
and fat-cells and forms high papillae, which are very well-developed, and the thickness reaches as high as 95% of the total thickness of the epithelium (in the case ofhumans oesophagus it is only 50%). The density of the papillae is also larger in the
cetacean oesophagus than in the human case.
The smooth muscle fibres of the lamina muscularis mucosae are arranged not soorderly as in other mammals, but are scattered irregularly. This unsystematicarrangement of the muscularis mucosae is noticed not only in the oesophagus but
also in every compartment of the stomach and throughout the intestine, thereby
losing the significance of the the name of " lamina ". The submucosa, formed byloose connective tissue, has nothing special to be described. The muscularis presents an inner, circular and an outer, longitudinal layer of muscle bundles. In theupper portion of the oesophagus it consists exclusively of striated fibres, but in the
middle and lower parts the muscularis is composed wholly of smooth fibres. Between the inner and outer layers there are scattered nerve fibres and ganglia, analogous
to Auerbach's plexus, which we come across in the intestines. The outer surface ofthe muscularis is covered with a thick membrane of connective tissue reinforced byelastic networks (adventitia).
The histology the first compartment of stomach is not very different from thatof the oesophagus, in spite of the slight difference in the relative thickness of thecorresponding layers or strata. It is quite obvious that this compartment isnothing but a dilated sack of the lowest part of the oesophagus.
The second compartment. The simple cylindrical epithelium lines the entire
inner surface of this compartment, including the gastric pits. It is composed oflight cells having distinct boundaries between each other. Nuclei of these cells
are small-sized, deeply stained and are situated at the base of the cells. At the floorof the gastric pits open numerous gastric glands occupying the propria and we candiscriminate two kinds of glands, the cardiac and the fundus glands.
The cardiac glands. Contrary to the earlier belief that the cetacean stomach is
destitute of the cardiac glands, we found cardiac glands to exist in the stomach of aSci. Rep. Whales Res. Inst.,
No. 23, 1971.

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HOSOKAWA AND KAMIYA

blue whale, occupying the narrow zone adjacent to the border-line at the junction
between its first and the second compartments. They are quite like the pyloric
glands of the third compartment in their structure, both being of the tubulo-alveolar
nature and composed of light cells of one kind.
The fundus glands. This type of gastric glands, branched tubular in nature,are extended in the greater part of the mucous membrane of the second compartment, situated within the propria and are 2-3 mm in thickness (in humans it is at
most about 1.0 mm). The zymogenic cells of this gland, 6-7 µ in diameter, arecharacteristic with the fairly coarse granules they contain, which stain with basic
dyes in the same way as the chromatin of the nucleus. The second kind of cells
found in this gland, the parietal cells, are much more developed than in the humanstomach. They differ from the chief cells in being considerably larger (10-20 µin diameter) and in their spherical or pyriform shape, containing sometimes two or
more nuclei. Frequently, especially in the lower part of the gland, they do notreach the lumen of the tube, being connected with it by a narrow pedicle. The moststriking characteristics of these cells are their strong affinity for acid dyes. Numeral
ratio between the zymogenic and the parietal cells is about 3: 1 and this index shows,compared with that of 5-6: 1 in the human case, that the cetacean stomach has
plenty of parietal cells, which are said to secrete hydrochloric acid. This peculiarityof the cetacean fundus glands must have some important meaning for the gastricfunction of this animal. Mucous neck cells are situated near the openings of theglands into the gastric pits and can be discriminated easily by their light cellularbody and somewhat shriveled nucleus, lying aside at the bottom of the cells.
The propria consists mainly of reticulated tissue, containing many free cells or
aggregated lymphatic cells.
The muscularis mucosae, which separates the propria from the underlying
submucosa, does not form a well-defined layer and is divided into the following fourgroups of muscle bundles; 1) muscle-fibres scattered in the propria, some of which
extend along the fundus glands up to the superficial layer underlying the epithelium,2) fibre-bundles running in the same direction as the reticulated folds of the mucous
membrane, 3) and 4) two muscle bundles in the deeper layer, which run independently of the direction of the reticulated folds, crossing each other at almost
right angles.
In the loose connective tissue of the submucosa run arteries provided with very
thick walls, just like the case in the cattle.
As muscularis was stripped of in our materials it was not possible to examine ithistologically.
The third compartment. The simple cylindrical epithelium clothes the inner surface, where numerous gastric pits are developed. Glands of this compartment that
extend into the propria are the pyloric glands. They are of the branched tubuloalveolar nature, and the glandular epithelium here presents great differences from thatof the fundus glands as these are composed of only one kind of cells. Althoughthese cells differ slightly from the zymogenic cells of the fund us glands, they resemble
the latter in all essential particulars so much so that we may take them for the same.
Sci. Rep. Whales Res. Inst.,
No. 23, 1971.

CETACEAN STOMACH AND FEEDING HABITS

97

The propria of this compartment does not differ from that of the second chamber. The muscularis mucosae also resembles that of the second one, but the firstgroup of muscle-bundles is more developed in this compartment and the second
group is lacking, while the third and fourth groups are only weakly developed.
In the submucosa there are scattered fat-cells, vessels, nerve-cells and fibres.
The muscularis is stripped off.
The fourth compartment (duodenal ampulla). The histology of the fourthcompartment in the oral half is not very different from that of the third compartment and in the anal half in their structure leads into the small intestine.

Stomach of the sperm whale
As refered to in the introduction, the constrictions between adjacent gastriccompartments in the sperm whale are marked more sharply than in the baleen
whales, and we can easily discern four compartments. The first and second compartments are very large, each measuring 140 x 140 cm for an adult male of 15.6 m
length, and the third and fourth are somewhat smaller, measuring 130 X 60 cm and
130 X 100 cm respectively.
The inner surface of the first compartment is covered with the same, yellowish
white epithelium as is the case with that of the oesophagus. There are, however,no well-developed folds, the inner surface here being provided only with the irregular
reticular fine furrows, especially in the region between the opening of the oesophagus
and the passage into the second compartment. The oesophageal epithelium of thefirst compartment stands out distinctly from the soft, mucous membrane of thesecond one. The surface features of the latter resemble that of the sei whale, having
been provided with the reticular folds, which diminish in the areas near the first
and third compartments. The third and fourth compartments are clothed by asimilar smooth membrane as in the sei whale, though in the fourth come into sight
semilunar folds as it approaches the opening into the small intestine.
Thickness of walls of different compartments. The thickness of the wall of thefirst compartment is 2.5-3.0 cm (mucosa 0.7, muscularis 1.7, serosa 0.4) of thesecond compartment 2.5-3.0 cm (mucosa 0.8, muscularis 1.4, serosa 0. 7), and eachof the third and fourth compartments 1.0-1.5 cm. It was not possible to do histological studies of the stomach of this whale, as it is very difficult to get seasonably
fresh materials to be used for microscopical work.
DISCUSSION

1.

Cetacean stomach seen from the comparative-anatomical viewpoint:

To understand the physiological meaning of the complicated feature of thecetacean stomach, we may first briefly review the comparative anatomy of thestomach in the animal kingdom. Fishes have stomachs of various shapes. Amphibians and reptiles have also gastric sacks and the birds posses in addition to thecrop, dilated sack of the oesophagus, glandular and muscular stomachs. As to themammals though the monotremata and a small group ofrodentia are said to have noSci. Rep. Whales Res. Inst.,

No. 23, 1971.

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HOSOKA WA AND KAMIYA

glandular stomach, almost all others have glandular stomach, where three distinct
regions or areas can be identified, namely the cardiac glandular, the gastric or
fundus glandular and pyloric glandular. Besides, there very often are also theoesophageal, non-glandular region or expansions. According to the combinationsof these regions among mammals, there are many types of stomachs, and Ellenberger(1926) classified them into the following groups:
1) Simple stomach composed merely of glandular regionsa) simple form lacking in cardiac sacksi) shaped like a simple sack, e.g. phoca;ii) provided with swelling of fundus, e.g. carnivora, insectivora,
many rodentia, chiroptera, apes and man;b) complex form provided with one or more cardiac blind sacks, e.g.
sirenian, large kangaroos and pig;2) Complicated stomach composed not only of glandular regions but also of
oesophageal region or expansions (always possess the cardiac area).
a) simple form ... the glandualr and oesophageal regions make altogether a simple stomacheal sack.
i) provided with not so large fundus, e.g. horse and tapir.
ii) provided with conspicuously large fundus, e.g. mouse and rat.
b) complex form ... the oesophageal ampulla makes one or more forestomachs independently of the glandular, real stomach, e.g. cetacean
and ruminantia.
From this classification it is clear that the complexity of the mammalian stomachs is caused mainly by two reasons by the expansion of the fundus or cardiac
blind sacks, and by the oesophageal forestomach that can be divided again into two
or more compartments. In view of these facts, the cetacean stomach, of which thecomplexity lies in the subdivisions of the glandular stomach itself, is an exceptionalcase in the animal kingdom. Though Ellenberger in his classification grouped thecetacean stomach with the ruminantia, we see now that there is a distinct difference
between these two. In the latter, though it is composed also of four compartments,
the first three, rumen, reticulum and omasum, are clothed with the oesophageal
epithelium and only the last one, abomasum, represents the real digestive part ofthe stomach. Thus in the ruminantia the stomach may be regarded as being primarily divided into two regions, the last of which only is the digestive region, and thefirst, non-digestive, being again sharply divided up into three compartments. In
the cetacea, on the other hand, the stomach, although divided primarily into twoparts like that of the ruminantia, shows a further subdivision of the digestive region
which may be exceedingly complicated in the Ziphiidae, while the non-digestive
region is not divided at all.

2. On the function of the cetacean stomach, with considerations on its interrelation to the foodcatching modus ef whales:
Anatomical features of various organs in the animal body are generally formed
and elaborated by two components, the external or the environmental and theSci. Rep. Whales Res. Inst.,
No. 23, 1971.

CETACEAN STOMACH AND FEEDING HABITS

99

internal or the hereditary. While the latter moment defines the hereditary features,the former modifies often the anatomical structures so as to be adapted to the environments of the animal. And such organs, under the influence of the environment, canget strongly modified in their shape, size and structure. We consider the stomach tofurnish an example of this latter kind, as this organ is apt to be modified according to
the alimentary life of its possessor.
From this view-point the complicated stomach of the cetaceans seems to be
intimately related to its alimentation, especially with its mode of food-catching in theaquatic environment. ·Weber (1885) mentioned already that the complexity of thestomach of the cetaceans was caused by the impossibility of their chewing in the aquatic
medium, which seemed to be quite acceptable to us. He, however, did not explain
the reason for the cetacean inability to masticate food. We would therefore, like to
put forward our view-point here to explain this.
Judging from the anatomy of the cetacean oral cavity, it is evident that thewhales do not masticate their food. To say nothing of the baleen of Mystacoceti, thenumerous teeth of Odontoceti are also far from being suited for mastication, for they
are conical homodont teeth, not to adapted for chewing, although aparently wellsuited for biting foods. The case of sperm whale and ,Ziphius also is not muchdifferent from that of Mystacoceti, as the former has only ten and several teeth in themandible and that with wide diastema in between the teeth, and the latter possess
only one or two pairs of teeth in the rostral end of the mandible. The mandibular
articulation and masticatory muscles of whales, together with the poor developmentof their lips also lend support to the above view, which is further augmented by thepoor development of the salivary glands indicating that the cetacean mouth is not
anatomically and physiologically meant for mastication. Thus deprived of thefunction of mastication, the cetacean mouth is devoted, we believe, exclusively toprehension of food.
Of the two major functions of mouth in higher animals, namely prehension andsalivary digestion, the cetacean mouth seems to be primarily concered with theformer function. This can be explained by the aquatic environment and foodhabit of these animals, as they require a large quantity of food, and like the ruminantia, they cannot be expected to waste their time between catching and swallowingof food. This feature needs a large stomach with compartments given to different
functions like storing, grinding and digestion. Thus the complex stomach of whales
with its four compartments is very well adapted for their aquatic life.
SUMMARY

Some of the notable findings on the anatomy of the cetacean stomach are summarized
as follows:
1. The cetacean stomach is composed of four serially arranged compartments,of which the first one is a oesophageal sack, the second one has numerous fundus
glands, the third one possesses many pyloric glands and the last one represents awell-developed duodenal ampulla.
Sci. Rep. Whales Res. Inst.,
No. 23, 1971.

100

HOSOKAWA AND KAMIYA

2. Between the second and the third compartments there is, moreover, asmall communicating division, which is prominent especially in the fetal stages.
3. The first compartment is dined with an extensively stratified fiat epithelium, of which the superficial layer is completely cornified and the cells have no
trace of nucleus, just like in the oesophagus.
4. The wall of the first compartment is the thickest of the four, and contains
very well-developed muscular strata. This compartment has undoubtedly a considerable contractibility.
5. Despite the general belief that the cetacean stomach is destitute of cardiac
glands, we found them in an adult blue whale stomach in the narrow region adjacentto the boundary between the first and the second compartments.
6. The fundus glands of the second compartment have extraordinarily numerous parietal cells. They may have some intimate relation with the gastric functionof whales.
7. The smooth muscle fibres of the lamina muscularis mucosae of all the four
compartments are scattered irregularly, and as such do not deserve to be called" lamina ".
ACKNOWLEDGEMENT
The authors are deeply indebted to Mr. K. Kureha of the Ocean ResearchInstitute, University of Tokyo for collaboration in collecting the baleen whale
materials.
REFERENCESBEDDARD, F. E., 1900. A book of whales. John Murray, New York. 320 pp.
ELLENBERGER, W. und H. BAuM., 1926. Handbuch der Vergleichenden anatomie der Haustiere. III Eingeweidelehre, 334-464, Springer, Berlin.
HUNTER, J., 1787. Observation on the structure and oeconomy of whales. Philosoph. Transact. Roy. Soc.
Lond., 16: 306-351.
JuNGKLAUS, F., 1898. Der Magen der Cetacean. Jen . ..(:'.F. Naturwiss., 32: 1-94.
SCHULTE, H. V. W., 1916. Anatomy of a foetus of Balaenoptera borealis. Mem. American Mus. Nat. Hist. New
series, 1 (VI): 389-502.
SLIJPER, E. J., 1962. Whales. 253-293, Hutchinson, London.
TYSON, E., 1680. Phocaena, on the anatomy ofporporse, dissected at Gresham colledge etc. Lond. (cited in
Jungklus, F., 1898)
WEBER, M., 1885. Anatomisches iiber Cetacean. Gegenbaur' Mikrosk. ]ahrb., 13: 616-663.

Sci. Rep. Whales Res. Inst.,
No. 23, 1971.

101

CETACEAN STOMACH AND FEEDING HABITSEXPLANATION OF PLATES
Plate I.
Mucous membrane of the stomach of a fin whale, 21.3 m long female.
Fig. I.
Fig. 2.
Fig. 3.

( X 3 natural size)

Boundary between the first (left) and the second compartments.
Boundary between the second (left) and the third compartments.
Boundary between the third (left) and the fourth compartments.

Plate II.
Histological section of the stomach of a fin whale, 21.3 m long female.
numerals showing the first to the fourth compartments. (x 4 natural size)
Fig. I.
Fig. 2.
Fig. 3.

Boundary between the first and the second compartments.
Boundary between the second and the third compartments.
Boundary between the third and the fourth compartments.
S: sphincter muscle

Sci. Rep. Whales Res. Inst.,
No. 23, 1.971.

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