THE.

SCIENIIfIC REPORIS
OF

Jff( Wllt\LES RESEARCH INSTITUTE
N0.3

THE WMLES RESEARQI INSTITUTETOKYO · JAPAN.FEBRUARY 1950

CONTaNTS
Page

1. On the Sensory the Tubercles of Lips and of Oral Cavity
in the Sei and the Fin Whale.
Teizo Ogawa, T; Shida· ...... 1

2. D:str.ibution of Red Marrow in the Bones of the Fin Whale.
Tadahiro Ohe·...... 17
3. On -the Cetacean Larynx, with Special Re~arks on 'th~ -;Laryngeal Sack of the Sei Whale and the ArytenoEpiglottideal of the Sperm Whale.
Hiroshi Hosokawa ...... 23
4. Bacteriological Studies on Freshness of Whale Meat.
(Repot No. 1)
Tomoichiro Akiba, Takeshi Tsuchiya,
Makoto Uemura. and Yoshiharu Natsume .. :' .. ·63
5. Protein Digestive Power of Sperm Whale.
Yasutaro Ishikawa······71
6. Properties of Fats and Oils contained in Various Parts of
Takajlro Mori, Masamichi Saiki ...... 79
a Sperm Whale Body.
. . .·
7. Studies on Kitol I.
Tadashi Tawara, Ryusuke Fukazawa···· .. 85
8. Studies on Kitol II.
Tadashi Tawara, Ryusuke Fukazawa ...... 39
9. Studies on Kitol III. Tadashi Tawara, Ryusuke Fukazawa··· .. ·92
10. On the Respiratory Pigment of Whale. (Studies on Whale
Blood II)
Tadashi Tawara···· .. 96
11. Research on Methionine in Whale.
Masami Yoshida .. · 102
12. Factory Ship Whaling around Bonin Island in 1948.
Kazuhiro Mizue··· 106
13. Statistic Study of Foetues of Whales.
Kazuhiro Mizue, Hisako Jimbo··· 119
14. Biological Survey of Fin and Blue Whales Taken in the
Antarctic Season 1947-48 by the Japanese Fleet.
Masaharu Nishiwaki, Kazuo Hayashi · · · 132'·'

Whales Reseach InstituteNo. 4, 12-chome, Nishigashi, Tsukishima, Chuoku
Tokyo, Japan
Copyright, 1950, by Meibundo Company
Printed in JapanAll rights in this reports are reserved.
Meibundo Company
No. 3, 1-chome, Kanda-Nishiki-cho, Chiyodaku, Tokyo, Japan

On. the Sensory Tubercles of Lips and of
Oral Cavity in the Sei and the Fin Whaleby
Teizo Ogawa and Toshitomo Shida
Department 'of Anatomy (Prof. Dr. T. Ogawa),
Medical School, University of Tokyo

Knowledges upon the sensory capacity of the cetacea are not only
important for the biological science, but must have also some practicalmeaning for the whalers. Recently a few works pertaining to this field
have been published from our laboratory, viz. upon the sinus-hair of theSei whale (Nakai and Shida), the auditory organ of the Fin whale (Yamada), and the intracerebral acoustic system of the whales (Ogawa andArifuku); the paper last mentioned contains the authors' summarized observations upon the serially sectioned brain-stems of Lagenorhynchus, Tursio,
Cogia, Physeter, and Balaenoptera.
In the study upon the sinus-hair of the Sei whale, Nakai and Shida were
induced to believe that the scanty but very regularly arranged hairs serve notmerely as a tactile organ in searching for food, but more probably as theorgan to feel the stream of water. They came to this assumption. chiefly
from findings on the inclination ·of the hair-shaft and the excentric positionof the hair-root within the venous sinus. Huber (1934) seemed also to haveconsidered that the sinus-hair of the baleen whales may afford a mechanism
for the appreciation of slight changes in water pressure.
While studying the sinus-hair,· one of us, Shida, directed his attention
to a large assembly of small granular tuberdes on the inner surface of
the upper and lower lip in the Sei whale, Balaenoptera borealis. The samestructure was seen, though less remarkably, not only on the palatal surface
sandwiched in between the right and left areas of baleens, but also on the
tongue and on the floor of the oral cavity. Microscopically examined, theycertainly represent highly sensitive tactile organs, performing no doubt afunction of much importance in the sensuous life of this whale. A little
later, we ascertaind in the Fin whale, Balaenoptera physalus, the existenceof structures approximate to those of the Sei whale. Though ;we have notyet compared other sort~ of baleen whales regarding this problem, it seems

2

Teizo Ogawa and Toshitomo Shid~ ·plausible that this sensory apparatus will be found also in other whales,
at. least in all members of the genus Balaenoptera.
1. Distribution of the granular tubercles on the lips and in the mouthcavity.

The tubercles in question are easily seen with naked eyes (fig. 8 a)on the inner surface of the lips as well as on the mucous membrane of
the mouth. The size of each tubercle and their number per unit area are
not equal according to the locality. On the upper lip., where their presenceis the most prominent, they are seen in a 16ng and broad zone, which
extends from the anterior extremity of the lip backwards to the ·angle ofmouth (fig. 1 a). The zone is of a breadth 3-4 cm at the anterior end,
6-7 cm at the middle part of the lip and becomes remarkably wider
towards the oral angle. There remains a narrow space of mucous membrane of a breadth about 1 cm, where the tubercles are wanting, between
the lateral margin of the bases of baleens and this zone (fig. 2) .. · At theanterior end of the upper lip the tubercles are relatively large, each ofthem measuring about 1 mm in diameter, in the hinder parts they aresmaller but more crowded, where it is difficult to determine their accurate
number. The density is ea. 200 per square centimeter at the anterior end(fig. 3), about 550 at the middle part of the upper lip, and nearly equal
or a little less at the posterior end.
On the lower lip (fig. 1 b), where in general the presence of the
tubercles is not so prominent as in the upper lip, they are the best seenin the posterior area near the oral angle, here about so clearly as in thehinder part of the upper lip. Forewards. they become within the lower lip
more indistinct, and at the most anterior portion the surface is everywhere
smooth, devoid of any remarkable tubercle, but· instead of this, many grayspots are visible on the surface, denoting the existence of well d,eveloped
papillae of the connective tissue underneath.
On the narrow palatal mucous membrane between the right and left
areas of baleens, relatively large granular prominences are present only inthe anterior part, about 1/5 of the entire length of the palate (fig. 1 a),while in the hinder part the surface looks quite smooth, though here also
we can recognize easily through the epitheli~m tops of many papillae ofthe connective- ti'ssue.

On the sensory tubercles of lips and oforal cavity in the Sei and tb,e Fin whale

3

Fig. 1. Jaws of Balaenoptera borealis; the granular tubercles are dotted.
sinus hair -- - ___ ---socalled
Jacobson's -- ~--- --organrows ofbaleens - - - -lower lip
(anterior part)-------·----upper lip(anteriorpart)floor of the -- ___ ,,...,,..,.•"'-mouth cavity

mucous,membrane of ~:::_
the palate--·tongue

upper lip
(posterior
part)lower lip
(posterior
part)(a) Upper jaw seen from below

Baleens(b)

Lower jaw seen from above

granular tubercles
assembled in a zone

I

''

/'
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_,---papillae of the connective
tissueFig. 2. A part of the upper lip of Balaenoptera borealis

4

Teizo Ogawa and Toshitomo Shida
Fig. 3. .Anterior end of th~ upper lips of Balaenoptera borealis
granular tubercles

socalled Jacobson's-- - -- - _____ _
organ
'

,'

baleensFig. 4. Balaenoptera borealisa) An excised part of the lingual surfaceb) Excised mucous membrane from the floor of the oral cavity

On the ·sensory tubercles of lips and oforal cavity in the Sei and the Fin whale5

In addition, the dorsal surface of the tongue and the floor of the oralcavity have also the granular tubercles nearly all over the surface, except
the most anterior parts (fig. 1 b), and backwards on the tongue they
The tubercles
become sparse, disappearing completely at the fauces.
disseminated in the posterior part of the oral floor do not show any clear

boundary against those in the hindermost portion of the lower lip~ On thedorsal surface of the tongue, where the mucous membrane forms a great
number of soft and gyrate folds, the tubercles are seen only on the summits, but not in the valleys between the folds (fig. 4 a).
The same
circumstance applies also to the floor of the oral cavity (fig. 4 b).
Inthese localities the density is about 70 per square centitneter in thenormally contracted mucous membrane. So when the membrane is distended, their number per unit .area will be much decreased.
In the Fin whale, compared with the Sei whale, we saw a differenceof little significance, only in regard to the upper lip, for the tubercles arewithin this lip the most easily recognized i:t1 the hindermost portion, andforewards tqey become relatively less distinct, being much smaller at theanterior end than those at the corresponding part of the Sei whale. But
as to the tubercles of the lo~er lip, the palate, the tongue, and otherparts of the oral cavity there is no remarkable difference between the twokinds of Balaenoptera.
2. Papillae of the connective tissue.
Microscopical examination reveals, that just under the tubercles papillaeof the connective tissue are very well developed, protruding deep ih thelayer of epithelium, and approaching very near the epithelial surface. The
form of these papillae is not always equal according to the locality, but
usually they are more or less swollen near the summit and show secondarybranchings. Especially at the anterior end of the upper lip of the Sei
whale the papillae are thick and high, providing with 3-5 elongatedbranches (fig. 5 a), while at the posterior part of the upper as well as .of
the lower lip they are in form somewhat like mushr~oms, having lessbranches (fig. 5 b). At the palate and at the anterior part of the lower
lip they are more cylidrical (fig. 5 c), and on the tongue as well as on thefloor of the oral cavity their form is irregular, though thick and short inmost cases (fig. 5 d).

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In the Fin whale we see very slender papiilae well developed, reachingnear the surface of the epithelium, at the anterior end of the upper lip,
wJ:iere as mentioned above the granular prominences are indistinct. As
to other places, no remarkable difference was ascertained between the twokinds of Balaenoptera, concerning the development of the papiIIae inquestion.
3. Nerve endings in and near the papiilae under the tubercles.
Within the papillae, especially near the summit of them we see many
terminal bodies of very characteristic form, to which sensory nerve ·fibers
are continuous. The intrapapillary terminal bodies are aggregated especiallyin a large number at the anterior end of the upper lip of the Sei whale.
But the apparatuses denoting the nerve ending are also found outside of
the papillae near the bases of them in the subepithelial layer. These
extrapapillary teqninal bodies are the most frequently seen at the anterior
end of both the upper and the lower jaws, less in other parts of the lips.
as weII as in the anterior portion of the palate. In these places we see here
and there within the depth of 0.4 min from the base of the epithelium asmall group of 2-6 terminal bodies, sectioned in various directions andaccompanied by a nerve bundle.
The gene~al form of the terminal bodies within or near the papillae
is known easily by examining the serial sections, stained simply withhaematoxylin-eosin. But we further tried block-staining by the Bielschowsky's method, and prepared the serial sections and stained them with van
Gieson's picrofuchsin, in order to study the finer structures, especially of
the axons.
In the basal part of the papilla the terminal bodies are comparatively
large in size, but less in number, while near the top of the papilla they
are relatively small but more numerous. (fig. 5). Calcul.ated at the base,each papiIIa seems to receive from one to several nerve bundles, each ofwhich contains a number of nerve fibers. Sometimes we see a very smalf
bundle, consisting of only a single nerve fiber. A teri:ninal body near the
top of the papilla measures 15-48 µ in breadth, while · its length can not
be easily determined, as its very long body is · wound in a complicatedmanner, not unlike the secretary part of a sweat gland. The terminal
body has in the interior a relatively thick, eosinbphile core and the core•

8

Teizo Ogawa and Toshitomo Shida

Fig. 6. Terminal bodies, not branched (a) and branched into three (b),both of which are much convoluted near the summit of a papilla (x500).
Reconstructed from sections.

knobs shortly before
the termination;
I
I~- terminal knobs_,
I.............. i

,,.
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.................................

' bifurcation
b)

',

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myelinated nerve fibers

is covered externally by several thin sheets of collagenic nature. Thisapparatus belongs certainly to the category of the ·lamellar corpuscles of
Vater and Pacini, but the number of lamellae is in this case very few.
So they seem to be the nearest to the Golgi-Maz~oni's corpuscles, as
described by Dogiel and van de Velde especially in the tactile balls of thecat. But compared with these corpuscles already known,. they are surpri-On the sensory tubercles of lips and oforal cavity in the Sei and the Fin whale

9

singly long and much convoluted (fig. 6). Because of the convolution asingle terminal body appears on the cut surface several times, which maylead to an estimation of them too much in number. But in reality,_ e~enat the anterior end of the upper lip,· where the terminal bodies are thebest developed, their number is in one papilla a little more than ten.
Each terminal body seems to receive only a single nerve fiber. But
we encountered bifurcations of the terminal body, the axon' with core andlamellae being ·divided into two or three equally thick branches, which areproximally united together, out have their ends separately (fig. 6 b). Sucha remarkable branching of the terminal body was observed especially at
the anterior end of the upper lip of the Sei whale. In the Fin whale
bifurcation of the axon occurs within the core very rarely.
The axon forms at the termination a small rounded or more irregularly
formed knob, which is not always the same in size (fig. 7). In the knob
we see a network or -a glomerule-like group of fine fibrils, with which
neurofibrils of the axon are directly continuous (fig. 7).
Moreover, we observed the axon in th~ terminal body a short distance
before its real ending to be divided suddenly into several fibrils ; the frayedfibrils are conglomerated, forming a knob, and then unite tOgether into asingle axon, and advance further to attain its real ending. Such a fraying
occurs 1-3 times during the course of an axon. Van de Velde seems
also to have seen th~ same relation in· the Golgi-Mazzoni's corpuscles of
the cat (1907, fig. 8 of his paper).
The extrapapillary terminal bodies in the subepithelial layer are elongated ovoidal or rod-shaped, provided with a core internally and with alamellated capsule externally. The number of lamellae is as few as within
the papilla. Also there they are wound, though the windings are not so
numerous, compared with the intrapapillary terminal bodies. They arevarious in size ; we measured a breadth of 38-170 µ and a length of 150-615 p (in the relatively straight ones). The core is 7-23 µ in breadth;this is swollen at the end usually to .a ·diameter of 30 µ. Within the corea single axis cylinder is contained, and its terminal portion is dilated intoa spherical or irregularly formed knob, with a network of fine fibrils.
4. Considerations upon the functional meaning.
The papillae mentioned above, being provided with so well developed

10

Teizo Ogawa and Toshitomo Shida

Fig. 7. Reconstructed figure, showing various
courses of nervous axons in a papilla.

) ~-' supE;rfici?l layer of-- the ep1thehum

deep layer of theepithelium

terminations of nerve fibers, are certainly an exquisite sensory organ forthe oral and labial stimulations. The impulses must be conducted fromthem to the brain-stem by the maxillary and . mandibular nerves of thetrigeminus.
Concerning the problem, whether the sensation caused· by these papillae
has anything to do with the gustatory sense, we suppose, that it can notOn the sensory tubercles of lips and oforal cavity in the Sei and the Fin· whale

11

belong in its nature to the sense of taste, for histologically the terminal
apparatuses, showing no intraepithelial nerve endings, quite different fromthe well known structures of the taste-buds. with the intra- and intergemmal nerve fibers. From every point of views this labial and oral apparatus
does not seem to be a chemoreceptor but merely a tangoreceptor, receiving
mechanical impulses, therefore very different from the taste.
On the other hand, we are at present uncertain, whether the cetacea
be endowed with the gustatory sense, for we have not yet found the taste. bud in them, though Wolf (1911) is said to have seen it in toothed whales.
Moreover, considering the meagre development or the complete absence ofthe olfactory nerves in the whales or dolphins respectively, it is quitepossible in our opinion, that the oral sense treated especially in the present
paper may substitute for the gustatory and olfactory senses in the cetacea.
It may also compensate in a certain degree the insufficiency of the visual
sense, for the comparatively small eyes of the whales, situated far behindand directed to the side, do not seem to be a very effective organ forthe perception of food entering the mouth.
The sinus-hair might also be effective as a tactile organ for food, butthey serve probably more for appreciation of the pressure or stream ofwater, having functionally much in common with the lateral line system of
the fishes. Nearly eighty years ago Malm (1866) reported an interesting
experience upon a Blue whale stranded alive at Goteborg, that " Der
Mann, welcher den Walfisch erlegte, bemerkte ebenfalls, dass die Lippen
unter den zuganglichen Teilen die empfindlichsten waren, da das Tierdurch Beriihrung derselben in die grosste Raserei versetzt wurde." Japha(1910) quoted this story to explain for the high sensitiveness of the hair for
touch, but in our opinion the granular tubercles with the richly innervatedpapillae are far more responsible for the sensitiveness of the lips than thehair. At present one of our eager wishes lies in a repeti~ion of such aninteresting experiment as Malm reported.
In the human anatomy it is well known, that the inner surfaces of thelips and of the cheek have in the newborn infants as well as in theadvanced foetuses numerous villi, which resemble therefore in localisation
the granular tubercles of the whales; they however begin to djsappear in
the .fourth week after the birth (Ramm, Heidsieck). In other mammals, it
deserves also noticing, that villi are. well developed on the mucous mem4

12

Teizo Ogawa and Toshitomo Shidabrane of the lips and of the cheek of ruminants and of many marsupials(Owen, Immisch, Schulze,· Sonntag). Though coincident in a certain degree
as to the localisation, the villi jn question of these animals seem to have
only a mechanical meaning in food-taking or in mastication ; they haveprobably not the meaning of highly sensitive organs as in Balaenoptera.
Neither can we expect any important function in the labial and buccal villiof the human being, which disappear very soon after the birth.
Lastly we wish to consider briefly the interrelationship between the
cetacea and other mammals. Since old time the opinion has prevailed, inthe realm of comparative anatomy, that the whales are the most kindred
to the ungulates. Relatively recently Anthony (1926) insisted upon theaffinity between the whales and the perissodactyla, and later Ommanney(1932), studying the urogenital system of the Fin whale, agreed with
Anthony on this point. On the other side, many palaeontologists havebeen inclined to look for the ancestor of the living cetacea in carnivora,
taking chiefly the dental characteristics of Zeuglodon in comparison.
Of course we can ourselves say nothing definite upon this important
problem, but would only refer to the fact, that the remarkable developmentof the palatal crests simultaneously with the high villous state of the labial
and buccal mucous membrane is met with in ruminants. And the baleensof the whales have been explained since Tullberg (1883) as spedally
differentiated sort of the palatal crests, an opinion, which seems to havemuch truth. Moreover, the palatal crests of some ruminants are closely
related to the baleens of whales, inasmuch as they are frayed at themargin. This combination of structures, that is, the simultaneous presenceof palatal crests and of labiobuccal villi, is found in a very accentuatedand specially differentiated form also in the whalebone whales. Naturally
it does not mean an important thesis in determining the relationship of thebaleen whales, but a comparative anatomical fact not unworthy of being
borne in mind at such a discussion~ Of course the toothed whales standhere out of the question, but we think ·there may be phylogetietically apretty large distance between the toothed and the baleen whales.
Summary.1. On the lips as well as in most parts of the oral cavity of the Sei whalea great many sensory tubercles are found, especially well developed at theOn the sensory tubercles of lips and oforal cavity in the Sei and the Fin whale

13

anterior portion of the upper lip. In the Fin whale the circumstance isnearly the same, with only the difference, that in this species the elevations
are at the anterior portion of the upper lip comparatively less distinct.
2. Microscopically examined, we see underneath the tubercles well developed papillae of the connective tissue, containing numerous sensory
apparatuses, with which the nerve fibers are directly continuous.
3. The terminal bodies in question show the structure the most approximate to the so-called Golgi-Mazzoni's corpuscles. But they are characterized
by the remarkable elongation. They have a core with the axon internallyand a few sheets of collagenic lamellae externally.
4. Su~cessive bifurcation of a terminal body into equally thick branches
occurs in the Sei whale. We found also sometimes the axon before its
real ending to be frayed out suddenly into a number of neurofibrils, which
after forming a knob unite together into a _single axon, to proceed further
toward its termination.
5. Because of the presence of the granular tubercles, the lips and the
mucous membrane of the oral cavity seem in Balaenoptera to be a highly
sensitive tactile organ, which is very important for the perception of foods,
while the sinus-hair are believed to have the functional meaning more forthe appreciation of pressure or stream of water.

Teizo Ogawa and Toshitomo Shida

14

Literature
R. Anthony:

Les affinites des Cetaces. Annales de l'Institut Oceanographique. ;Nouvelle serie.

3, 1926.
A. S. Dogiel : Der fibrillare Bau der Nervenendapparate in der Haut des Menschen und der Sau·
getiere und die Neuronentheorie. Anat. Anz. 27, 1905, 97-118.
K. B. Immisch:

Untersuchungen iiber die mechanisch·wirkenden Papillen der MundhOhle derHaussaugetiere. Anat. Hefte. 35, 1908, 759-759.
A. Japha: Die Haare der Waltiere. Zoo!. Jahrb., Abt. f. Anat. ;n. (Habilitationsschrift, 1910, 143).
E. Heidsieck:

Die Ruckbildung der menschlichen Lippen- und Wangenzotten.

Anat. Anz. 61,

1926, 321-347.
Ernst Huber: Anatomical Notes on Pinnipedia and Cetacea.

Carnegie Inst. Wash. Pub!. 447,

1934, 105-136.
A. W. Malm:

Einiges von den Walfischen im Allgemeinen und Balaenoptera Carolinae im

Besonderen. Berlin, 1866. (Cited from Japha, 1910, p. 36-37).
J. Nakai and T. Shida: Sinus hairs of the Sei-whale. Scientific Reports of the Whales ResearchInstitut, Tokyo. 1, 1948, 41-47.
T. Ogawa and S. Arifuku:

On the Acoustic System in the Cetacean Brains. Idem. 2, 1949, 1-20.

F. D. Ommanney : The Urino-genital System of the Fin Whale.

Discovery Reports. 5, 1932,

363-466.
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II. Beuteltiergattung Macropus (Shaw). Idem. 1913.
T. Tullberg: Bau und Entwicklung der Barten bei Balaenoptera Sibbaldii. Nova Acta Societatis
scientiarum Upsala. (Cited from w. Leche in Bronn's Klassen .und Ordnungen. 6, Abt. 5,
1887-1898, and from G. Retzius in Biol. Unters. N. F. 13, No. 15, 1906, p. 122).
Em. van de Velde: Die fibrillare Struktur in den Nervenendorganen der Vogel und der Saugetiere. Anat. Anz. 31, 1907, 621-634.
E. A. Wolf:

Ueber die Zungenpapillen der Wale. (Dissertation) Berlin, 1911. (Cited from S. Schu-macher in Mollendorff's Handb. d. mikrosk. Anat. 5, Tei! 1, 1927, p. 56).
M. Yamada : Auditory Organ of the Whalebone Whales. Scientific Reports of the Whales ResearchInstitute, Tokyo. 2, 1949, 21-30.

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Distribution of the Red l\!Iarrow m Bones of the Fin \tVhale
by
Tadahiro Ohe
Department of Anatomy (Prof. Dr. T. Ogawa),
Medical School, University of Tokyo.

The present investigation was performed at Mombetu, Hokkaido, inthe summer of 1948, when I stayed there for ten days. The object of mystudy was to determine the site of haematopoesis in the body of whales,
for then we had in mind a question, whether or not the blood corpusclesof the baleen whales are developed in the organs just as they are in
the terrestrial mammals. Except the quite recent paper of Slijper et al(1948), which reached us after the completion of the present work, there
seems to have been no detailed report upow the distribution of the red bonemarrow in the baleen whales.
Materials:.
Four individuals of the Fin whale, Balaenoptera Physalus L., were
examined.
All of them were captured off the northeastern coast of
Hokkaido by boats of the Taiyo Fishering Company ; by the courtesy anddeep understanding of the members of this company the present work
could be fulfilled to the success of a certain grade.
Methods:
Of course at the whaling station the scientific research is met withmuch difficulties, as speedy disposition of the whale body is the mostimportant factor in the whaling industry and in most cases the heavymaterials can only be moved by the help of winches. We can examine
them not always to our hearts' content.
My own experience at thatoccasion is as follows. The vertebral column, .after most of the truncal
muscles were cut off, is brought by the winch to the "bone ·platform";the huge mass was separated into each segment, by cutting through theintervertebral discs. At that time I endeavoured to arrange all of thevertebral bones in the natural order, and asked one of the workers to
break them lengthwise in the oro-caudal direction. Of some vertebrae thetransverse and the spinous process were also cut for examination of thebone marrow. The cervical vertebrae were split longitudinally in a mass.

Tadahiro Ohe

18

The dissectors used to remove the ribs altogether with intercostal muscles
and pleura as a whole from the vertebral column, then to separate theminto each rib and to bring them to the bone platform. At that occasion I
arranged the ribs in the natural order on each side and examined theircaput, cauda and the middle part, sectioning transversely. Though theribs of the Fin whale are said to be usually fifteen pairs, there were twocases Nos. 42 and 40, in which I found only fourteen pairs. The problem,
whether there were really only fourteen pairs, or one pair, probably thelast, was lost at the dissection platform, is not certain. I guess the former
assumption may have more truth.
To my regret, I had no chance to examine sternum and pelvis. I did
not dare to find out these small bones, not to disturb too much the speedy
work at the dissection platform. The chevron bones are carried to thebone platform as a long chain-like block, and I cut some of them to see
the marrow.
The large skull is at first divided by the bone saw lengthwise at the
middle line into right and left halves, cut then in various directions into
many blocks, which are small enough to be easily dropped into the pressboiler. I examined the cut surfaces of them as much as possible. The hyoidconsists of one median part (basihyal and thyroyal) and paired lateralparts (stylohyal). The stylohyal was usually not discovered, as it was
carried probably to another place. I cut as a rule .the hyoid bone in the
median plane and further along its long axis.
The scapula was cut into two parts, a cranial and a caudal. The large
projected acromion was examined especially. Bones of the pectoral limb
were cut in a mass transversely into two pieces, and then in the same
direction into four portions. The head of the humerus was especially·observed.
I made some microscopical preparations from the red marrow andascertained· that the haematopoesis was certainly taking place in it.
Results:
No. of the specimen----- ----~

- - - - - - - --------

44

!

41

40

I

42

50

I

52

53

I

67

Male

FemaleFemale5 p. m.
Augu!;t 12.

5 p. m.
August g.

8 a. m.
August 12.

------

Body length in feet------------------

Sex

FemaleDate of the capture

8 a. m.
August 14.

!

Distribution of the red marrow in bones of the· Fin whale

Beginning of dissection

6 p. m.
August 14.

Approximate hours elap·
sed between captureand dissection
Vertebral formula<U<U

6 a. m.
August 10.

10

10

11

7+15+14+20=50

7+15+14+22=58

4 P· m.
August 12.

I

I

8

Not especially examined.

Atlas

The ·red marrow was found only in the ventral arch.

Cervical vertebrae 2- 7

Entirely filled with red marrow, but in No. 44 not so completely as in three other individuals.

Thoracic vertebrae
1-15

Also filled with red marrow, but the white marrowCl) was present in a wedge-like formC2l in the peripheral I part of thevertebral body, becoming larger in the caudal direction.C 3X 4)..a"'.......
>I Lumbar and caudal<U

3 a. m.
August 13.

I

I

19

Beginning with the 15. (in No. 42 with 14.) thoracic vertebra
caudalwards the red marrow becomes irregular in shape andgradually decreases in extent, but in Nos. 41 and 40 it waspresent in addition along the lines of epiphysis. A red spot
vertebrae
was found down to the 2. caudal (No. 44), to the 12. lumbar(No. 41), to the 8. lumbar (No. 40), or to the 2. caudal (No.
I
42).
--:Processes-- examinecf
6. thor.+10. thor.+
5. thor.+~ tho<.+"'
of some vertebrae ( + :15. thor.14. thor.7.
thor.+ I 10. thor.+
I. lumb.f
presence of the red mar·
10. thor.+row;-: its absence)
14. thor.One red por- ; Though I reTwo small
The
skull
pieces of red tion was pre- member to ha- had the redmarrow, each sent in the ba- ve seen the red marrow only
the size of a sal part and ' marrow in the in its anterior
hen's egg, we- another in the i skull, I can not basal part.
Skull
re present in frontwall of the i clearly locate
the basal part, brain case and · it.
and a larger one more in theI
one in the roof occipital part.
of the brain case.

I

1-

I

Not
Hyoid
examined.

A . relatively : The red marlarge red por- row was pretion in the thy- sent only inrohyaland the thyrohyal.
scattered red The stylohyal
spots in the. was not obserbasihyal.
ved.
Stylohyal note}l1amined.

The red marrow was present neither inthe basi- andthyrohyal, nor
in the sty lohyal.

I
Ribs
Glenoid

"'
'3p.
r.n"'

Scapular plateu

The red marrow was present only in the caput. But only inthe caput of the 10. rib of No. 41, the red marrow was mixed
with the white one.
The red spat was seen.
Three veryHereandsmall spots of there red marthe red marrow row was prewere found.
sent.

Acromion

No red marrow

Bones of flipper

No red marrow

No red m'arrow

20

Tadahiro Ohe

Supplementary remarks to the table of the forego'.ng pages :(1) Man says usually "red" and "yellow" as to the bone marrow, but
in the whale, especially in the vertebrae of them, the yellow marrowmight be better called ''white", because in the lumbar vertebrae, forinstance, we get an impression, as if red ink were dropped on the snow.
So I used in this paper, instead of " yellow ", the term " white bonemarrow." But in hyoid, scapula, and bones of the skull the yellowish
colour predominates.
(2) By cutting further, the red marrow was seen in form of an hour-glass.
(3) In the epiphyses of the thoracic vertebrae of No. 42 the red marrowof a patch-like form is present in the center, surrounded by the whitemarrow ; this fact was not seen in the other specimens.
(4) In the 10. to 13. thoracic vertebra of No. 42 the dorsal wedge of thewhite marrow was relatively sharp and small, while the ventral whitepart showed more rounded contour.
(5) The red marrow .in the processes of the 4. and 7. thoracic vertebra
was continued to that of the vertebral body, while that in the processesof the 10. thoracic vertebra showed no direct continuation with that of
corpus.
Summary:(1) In the atlas the red marrow is present only in the ventral arch. Fromthe 2. cervical down to the 14. thoracic vertebra the vertebral bodies arefilled nearly completely with the red marrow, and as the whale advancesin age, this seems to disappear in the caudo-cranial direction.
Beginning with the 15. thoracic verte4ra the red marrow becomes
irregular in shape and decreases suddenly in extent, but point-like redplaces can be found up to lumbar ·or caudal vertebrae.
As Slijper says, the red marrow seems to disappear earlier in theprocesses than in the corpus of vertebra. In the latter the white ·marrowappears at first in a wedge-like form in the circumferential part, and atthat time the convertion of the red marrow into the white has not yet
finished in the processes.
In the vertebrae of two individuals (No. 44 and No. 41), the redmarrow, irregular in shape, was also present along the lines of epiphysis,
but this wai;; not the case in the other two (No. 40 and 42).
(2) In scapula, hyoid and skull the red marrow becomes smaller with theDistribution of the red marrow in bones of the Fin whale

21

age. Slijper et al seem in the Blue whale not to have seen the red marrow in these bones.
(3) In all of the costae the red marrow was present in the caput.
(4) In bones of the flipper and in the chevrons the red marrow is completely lacking.
Explanation of the figures :Except hyoid, lateral views of all of the bones are shown. The processes of vertebrae areomitted ; ribs, hyoid and bones of the flipper are moved from their natural positions.
black·························································red bone marrow.
obliquely lined area···· · · · · · · · · · · · · · · · · · · · · · · · · · · ······bones not examined.

Literature :C. F. Feltmann, E. J. Slijper and w. Vervoort: Preliminary researches on the fat-content ofmeat and bone of Blue and Fin whales. Koninklijke Nede.rlandsche Akademie van Weten·
schappen. Vol. 51, No. 5 (1948) p. 604-615.

Tadahit'O oheOn the Cetacean Larynx, with Special Remarks on the
Laryngeal Sack of the Sei \Vhale and the ArytenoEpiglottideal Tube of the Sperm vVhale
byHiroshi Hosokawa
Department of Anatomy (Professor Dr. T. Ogawa),
Medical School, Universityof Tokyo.

Introduction
Since very early the larynx of the whales has, because of its peculiar
structure, attracted notice of many anatomists. The most striking peculiarities lies in the tube formed by the upward elongated epiglottic and arytenoid cartilages and dilated at its summit like a knob, protruding into thechoana. After Tyson (1680), who observed this tube for the first time in
Phocaena communis, a number of anatomists described this structure invarious kinds of whales, for instance, J. Hunter (1787) in the harbour
porpoise, dolphin, pottfisch and narwhal, Meckel (1833) in Delphinus andPhocaena, Stannius (1846) in Delphinus and Monodon, Vrolick (1848) inHyperoodon, Huxley (cited from Howes), Burmeister (1867), Howes (1879),
Rawitz (1900) and Boenninghaus (1902) in Phocaena communis, Murie(1871, 74) in Grampus rissoanus and Globicephalus, Watson and Young(1878-79) in Beluga leucas, Dubois (1886) in Hyderoodon, Tursiops, Phocaena, Delphinus and Globicephalus, and Benham (1901) in Cogia breviceps.
To the last mentioned author we owe probably the name " arytenoepiglottideal tube ".
Meanwhile it is worthy of notice that research materials of theseanatomists were always toothed whq.les. The question naturally arises,
whether the whalebone-whales have also the same tube in the larynx..
This problem was answered by Eugen Dubois (1886), who compared
the larynges of Hyperoodon, Tursiops, Phocaena, Delphinus and Globicephalus with that of a fetus of Balaenoptera Sibbaldii, and asserted thenon-existence of the aryteno·epiglottideal tube in the latter. Besides there
are, according to him, m1ny other differences in the larynx between
whalebone and toothed whales.
While engaging recently in the anatomical research up.on some whales,

24

Hiroshi Hosokawa

I was much interested by the striking difference in the larynx between
two groups of the whales and was led to make some detailed inquiries
about this matter.
The following . materials have been hitherto examined by me in thisresearch. Whalebone whales :1. Balaenoptera borealis, 42 feet, female; 46 feet, female (at Ayukawa,
Miyagi Prefecture).
2. Balaenoptera musculus, fetuses of various lengths (in Antarctic
Ocean).
3. Balaenoptera physalus, fetuses of various lengths (in Antarctic Ocean).
Toothed whales:·
4. Physeter catodon, 32 feet, male; 36 feet, female (at Ayukawa).
5. Lagenorhynx obliquidens, male fetus of 51 cm length (at Kawana,
Shizuoka Prefecture).
6. Prodelphinus caeruleo-albus, 11 feet, male; 117 cm, female (atKawana).
7. Globicephalus melas, 15 feet, male; 16 feet, female (at Ayukawa).

In this1 paper I will treat mainly the larynx of the Sei whale (Balaenoptera borealis) and that of the Sperm whale (Physeter catodon); observations on the larynges of the others will be reported later.
I. Cartilages of the larynx
The larynx of the whales is constructed, as in other mammals, of the
cartilagenous framework and several muscles connecting the cartilages.
They form as a whole a tubular organ with the laryngeal cavity in it.
The inner surface is covered with the mucous membrane continuous withthat of the pharynx upwards and of the trachea downwards.
The laryngeal cartilages are five in number, three of which (thyroid,

cricoid, epiglottic) are unpaired and the rest two are paired (arytenoid). The
corniculate and the cuneiform cartilages do not exist as separated ones ..
At first the form of each cartilage will be mentioned, compared between
the Sei and the Sperm whale.
The THYROID cartilage
Sei whale: (fig. 1) Two laminae, right and left, of this cartilage arefused together iq_ the anterior middle line, there forming no sharp angle.

On the Cetacean Larynx

25

The cartilage is as a whole a plate concave behind and shaped like a
flying swallow, providing with four processes; the inferior corun, corresponding to wings of the swallow, is very well developed, projects caudally andmakes a joint with the cricoid. - The upwards directed superior cornu is
not so well developed. Between the superior horns of both sides a deep
V-shaped notch exists, separating partly the right and the left lamina(incisura thyreoidea cranialis). The caudal border has in the median line
a shallow notch (incisura thyreoidea caudalis).
The outer surface of the laminae, smooth and convex as a whole,shows no remarkable prominence. The thyreohyoid and sternothyreoid
muscles are attached to relatively small areas near the superior lateralborder of the lamina. The cricoarytenoid muscle inserts partly to theouter surface around the caudal notch, but for the greater part to theinner surface of this cartilage. To the inner surface are attached also
three muscles, thyreopharyngicus, thyi:eoarytaenoideus and thyreoepiglotticus. Moreover the posterior end of the inferior horr1° is connected withthe musculus cerato-cricoarytaenoideus dorsalis.
Sperm whale: (fig. 1') The thyroid is the largest cartilage in thelarynx of this whale, differing in shape very little from the homologous
structure in most of other mammals. Two laminae, right and left, arefused together at the middle front, forming a tolerably sharp angle, andthere we see the prominentia laryngica.
The superior horn is very slight, and .the inferior one is also not sowell developed as in. the Sei whale. Each lamina is irregularly triangularin shape ; its outer surface is not smooth, having along its upper border alineal crest, which courses downward to the middle portion of the lamina,
and is divided there in two crests ; one of them goes to the dorsal side,slightly inclining to the caudal direction, while the other runs ventrocaudally, and serves as insertion for the thyreohyoid and sternothyreoid
muscles. At the point, where the lineal crest changes its course from theupper border to the middle of the lamina, it surrounds a pore, foramen
thyreoideum, through which a nerve (the inner branch of then. laryngicus.
cranialis) and a vessel (a. laryngica cranialis) penetrate. Existence of sucha pore was mentioned already by Stannius in various whales, and by
Rawitz in Phocaena communis.
The inner surface of the lamina is nearly smooth and gives attachment

26

Hiroshi Hosokawa ·to the thyreopharyngeal and thyreoarytenoid muscles etc.
The EPIGLOTTic cartilage
Sei whale : (fig. 2) This elastic cartilage is not so hard as other larygeal
cartilages. Its upper half, not so dilated widely as in the human larynx,
is concave behind and convex in front. The lower half (petiolus epiglottidis) is long and thin, and directed to the cranial notch of the thyroid
cartilage.
Sperm whale: (fig. 2') This large cartilage is an elongated triangle
in shape seen from lateral, with an apex directed ventro··cranially, where
the hyo-epiglottic muscle is attached. The cranial apex is, seen from behind,triangular arid involved forwards.
The hind surface of this cartilage has three lineal longitudinal elevations, one in the median line and two others along the lateral margins ; thelatter meet with the lateral border of tihe arytenoid cartilage and areconnected with it by the mucous membrane, forming so together thea:r.yteno-epiglottideal tube.
The caudal end of this cartilage has a tubercle, whioh is connected tothe angle between two laminae of the thyroid cartilage, a little below thecranial noch. This connection is strengthened by soild ligamentous fibres.
The Cmcom cartilage
Seiwhale: (fig. 3) The name "cricoid" does not hold good for thiswhale, for the cricoid cartilage is not shaped lik;e a signet-ring as in othermammals, but its ventral half, corresponding to arcus, is completely deficient and the whole cartilage is merely a plate concave in .front. The
upper margin of this plate has a slight prominence at the middle, and alittle more laterad on each side an articular facet for the arytenoid cartilage.
Caudally it is directly united with the tracheal cartilages, forming
together the cricotracheal plate in the dorsal wall of larynx and trachea.
Lateral parts of this plate are partially separated by a longitudinal notchfrom the main medial portion and may be named " processus lateralis" ;the inner concave surface of this lateral process gives attachment to a
portion of muscles of the laryngeal sack, which will be mentioned later.
Moreover, the m. cricothyreoideus inserts to the outer surface of thisprocess, a little above its caudal end.'

On the Cetacean Larynx

27

Inner and outer surfaces of the cricoid cartilage, nearly smooth all over,have a slight crest along the median part of the outer surface, which runs
from the upper border downwards, remarkably dilating. Wide areas on
both sides of the crest are for insertion of the dorsal cricoarytenoid
muscles. The most caudal part of this area is connected with the criocopharyngeal muscle. A little more mediad than the upper end of the notch,
which demarcates the lateral process, there are artiCular facets for theinferior horns of the thyroid cartilage. This joint is not fully diarthrodial
but somewhat syndesmodial, strengthened behind by fibrous bands, while
the aryteno-cricoidal joint is diarthrodial, provided with the articular capsule.
Sperm whale: (fig. 3')
This whale has a completely ring-formed
cricoid cartilage, a very diferent circumstance therefore from the Sei whale.
This fact deserves special attention, for, according to Dubois, all of the
toothed whales· he examined, with the exception of Tursiops! have incompletely ring-formed cricoid cartilage ; in them the ventral portion is said tohave a fissure, though the fissure is not so broad as in whalebone whales.
Sanctis (1879) seems to have found in Physeter macrocephalus the same
matter as in Tursiops (cited from Dµbois); and Benham found also in an
adult male Cogia a completely ring-shaped 1cricoid catilage. In other toothed
whales examined by me the matter is quite the same as Dubois described~
Therefore, as to the cricoid cartilage there are ·two groups in the toothed
whales, one of which (Physeteridae etc.) has a completely ring-shaped
cricoid cartilage, while the other (Delphinidae and Hyperoodon) an incomplete one. The cricoid cartilage of the sperm whale is not united withthe tracheal cartilages, a distinct space filled with fibrous bands existing
between them.
The very thick posterior portion of .the cricoid cartilage gives attaehment to the dorsal cricoarytenoid muscle, while the middle portion of its
cranial border is connected with some fibres of the transverse arytenoid
muscle. The anterior part of this cartilage is narrow and somewhat thin,
but widens as it approaches the median line.
The ARYTENOID cartilages
Sei whale: (fig. 4) Each of the paired cartilages has two· processes
directed up and down (processus cranialis and proc. caudalis). The tongue·
shaped cranial one is thin in the transverse direction, while the caudal one

28

Hiroshi Hosokawa

is dick and rounded. The pointed ends of the caudal processes are connected by fibrous tissue between both sides. The main part of the cartilage,
corpus, has a laterally directed large prominence, which corresponds
probably to the processus muscularis, giving attachment for the dorsal
cricoarytenoid muscle etc. The dorsal surface of this portion has an
oblique oval facet to articulate with the upper border of the cricoid
cartilage. The corpus has furthermore a crest along the medial border ofits ventral surface, probably homologous to the processus vocalis. · Uponthe relation of various muscles to this cartilage 1 want to refer it to theaccompanying figures.
Sperm whale: (fig. 4')
The most remarkable difference upon thearytenoid cartilage between Sperm and Sei whale is the absence of caudal
process in the former. In the Sperm whale the well developed cranial
processes of both sides, standing side by side on the cricoid cartilage, form
a concave furrow in front, and their upper ends are fused together into one
cartilage. Between the ventrolateral border of this process and the lateral
crest of the epiglottic cartilage a fold of mucous membrane (plica aryepiglottica) is stretched and thus the aryteno-epoglottideal tube is completed.
The CoRNICULATE and the CUNEIFORM cartilages
These cartilages could not be found in all the whales I have examined,
at least as separated ones. Almost all of the authors who have studied thelarynx of whales did not mention about them. Watson and Young denied
the existence of them in ·Beluga. But Howes remarked in a fetus of
Phocaena that the arytenoidi cartilage consists of two separated pieces,and the 'upper portiop. of the so-called arytenoid, which is prolonged into
the nasal passage, must represent an elongated cuneiform cartilage, theSantorinian element being absent. Negus (1928) agreed with Howes; he had
examined a reconstruction model of the larynx of a fetal Sperm whale,
which was 4.5 inches long, and finding two columns of mesenchymal,
cartilaginous cells lying above and slightly ventral to the arytenoids,
mentioned confidently· that these cell-groups would make the ca1tilages of
Wrisberg and, together with the epiglottis, constitute the peculiar laryngeal
tub.e. Of the Santorinian cartilage he denied its existence. In sm3.ll fetusesof Blue and Fin whales I examined minutely the cranial process of thearytaenoid, but no separated piece of the cartilage was found, and in a

On the Cetacean Larynx

29very small fetus (17 cm of length) of Megaptera nodosa, I saw that only the
corpus of the arytenoid consists of cartilaginous element, the cranial process
being formed merely of connective tissue.
The TRACHEAL cartilages
As these are closely related to the larynx, I will give a brief description of them.
Sei whale: (fig. 3) All of the tracheal cartilages are continuous witheach other and moreover, as mentioned above, with the cricoid cartilage intheir dorsal parts, forming altogether a single plate. In the lateral parts
they are however separated from each other by tranverse ·fissures and the·ventral portions are completely wanting. Though Dubois stated that only
the upper 4:-5 tracheal cartilages are fused with the cricoid in a fetus, of
B. Sibbaldii, in my observation of the Sei whale all of the eight cartilages
up to the bifurcatio trancheae form a single plate.
Sperm whale: (fig. 5') In this whale the tracheal cartilages are complete rings, and the i;nost cranial one is separate from the cricoid cartilage.
The upper three cartilages are fused together in the dorsal part~, but theothers are all separated from each other. The fact, that all of the tracheal
cartilages form complete rings, seems to be an exception in the whales.
For, according to Dubois,. the uppermost one (Hyperoodon) or two (Delphinus, Phocaena) tracheal cartilages are lacking of its ventral portion,
and also Watson and Young mentioned the same relation as to two or
three upper cartilages respectively for Beluga and Delphinus.
II. ' Muscles of the larynx
Laryngeal muscles of the Cetacea have been studied by a number ofanatomists, but their explanations are not always coincident. The differenceof opinions come, to my mind, partly from difference of the materials
treated. In fact, every species of the whales has more or less its own
structure. We must naturally consider seriously the differences between
toothed and whalebone whales. In the following my results upon laryngeal
muscles of the Sei and the Sperm whale will be mentioned.
Sei whale:
As in the case of other mammals, the laryngeal muscles are classified
also in this whale into two groups, extrinsic and intrinsic. The· extrinsic

30

Hiroshi Hosokawa

muscles, connecting the larynx with neighbouring structures, are :Musculus sternothyreoideus (innervated by branches from the ansa
hypoglossi)
Musculus thyreohsi-oideus (ditto)
Musculus laryngopharyngicus (innervated .by branches of the plexus
pharyngicus)
Musculus hyoepiglcttious or baseohyoepiglotticus
The STERNOTHYREOIDEUS and the THYREOHYom:Eus belong to the so-called
infrahyoidal muscles and insert to the thyroid cartilage. The well developed
LARYNGOPHARYNGICUS consists of THYREO}'HARYNGICUS and CRICOPIIARYN·a1cus. The former arises from the frontolateral part of the inner surfaceof the thyroid cartilage and its massive fibres surround the pharynx tomeet with that of the opposite at the posterior median part of thepharyngeal wall, i. e. raphe pharyngis. The latter originates from thedorsolateral part .of the cricoid cartilage and, running up and medial wards,
terminates also at the raphe pharyngis. The unpaired HYOEPIGLOTTrcus
was described by Carte and Macalister (1868) in Balaenoptera rostrata, but
I could not see it in the Sei whale.
The intrinsic muscles are those, which begin and end within the larynx
itself. In the following I will describe them one by one.
( 1 ) M. CRrcOTHYREOIDEus; (fig. 5) This muscle arises from the outersurface of the lateral process of the cricoid cartilage and runs upward
and laterad, to insert on the medial border of the inferior horn of the
thyroid cartilage. It is not divided into two parts, pars recta and parsobiqua, as in the human larynx. This muscle is the only one in the wholelarynx, that is innervated by the cranial laryngeal nerve (fig. 18).
( 2) M. CRrcoARYTAENOIDEUs DoRSALrs; (figs. 6, 7) This muscle arises
mostly. from the posterior medial portion of the cricoid cartilage, partiallyalso from the caudal end of the inferior horn o~ the thyroid cartilage. Its
fibres converge upwards and laterad to reach the muscular process of thearytenoid cartilage. Because of the connection with the inferior horn, it
may be called better " m. ceratocricoarytaenoideus dorsalis ", the nameused already by Dubois.
( 3) M. ARYTAENOIDEus; (fig. 16) Tracing the fibres of m. cricoarytaenoideus dorsalis upwards, we come across muscle fibres running transversally between the arytenoid cartilages of both sides. The fibres ariseOn the Cetacean Larynx

31

f~om the dorsolateral margin of the cranial process of the arytenoid cartilage on one side and attain the corresponding part of the same cartilageof the other side. This muscle (m. arytaenoideus transversus of Dubois)
corresponds probably to the transversal part of m. arytaenoideus in thehuman larynx. The pars obliqua is not found in the cetacean larynx.
According to Dubois the most caudal part of the arytenoidal muscle iscovered by the upper part of the dorsal cricoarytenoid muscle, and moreover some fibres of the former are attached to the cricoid cartilage. On
this point I agree with Dubois, though the boundary between these two
muscles is not very distinct.
( 4 ) Muscle of the walls of the laryngeal sack (M. THYREOARYTAENOIDEUS,
M. CRICOARYTAENOIDEUS LATERAUS, M. ARYEPIGLOTTICUS, M. THYREOEPIGLOT'l'ICUS ) ; (figs. 5, 8, 9, 12-17)
As will be mentioned later, the Sei whale
has a large sack attached to the ventral wall of the laryngotrachea. It
communicates with the proper laryngeal cavity through a wide opening
which exists between the arytenoid cartilages of both sides. This laryngeal
sack has thick muscular walls and its muscles, cross-striated in nature,
are arranged so differently from the laryngeal muscles in other animals,that it is not easy to determine the homologies between them.

Carte and Macalister examined for the first time these muscles in
Balaenoptera rostrata with the conclusion that they belong to m. thyreoarytaenoideus. Dubois insisted that not only the thyreoarytenoid muscle
but also the m. cricoa:rytaenoideus lateralis takes part in the formation ofthis muscular wall. Thereafter Benham, studying a newborn female Balaenoptera rostrata, supported Carte and Macalister's opinion and denied theco-existence of the lateral cricoarytenoid muscle.
At any rate these muscles exhibit quite strange arrangem~nt in relation to the cartilages. In the Sei whale we see on the ventral surface ofthe larynx them. cricothyreoideus on both sides. Moreover,- massive muscle
fibres come into sight from behind the caudal ridge of the thyroid cartilage,
nmning downwards and ·being directed in the lateral parts somewaht
laterad. These muscles, which form the ventral and lateral walls of thelaryngeal sack, insert partly to the ventral ridges of the cricoid andtracheal cartilages, where these cartilages are defective in the ventral
portion. More cephalic and deep fibres of them reach the muscular processof the arytenoid cartilage.

32

Hiroshi Hosokawa

The comparatively superficial muscular layer of the laryngeal sackcourses as a whole in the caudo-lateral direction ; I would call it "A-group"in the following description.
A deeper group of muscle fibres arises mainly from the ventral surfaceof the caudal process of the arytenoid cartilage and, diverging downwards,
forms a part of the ventral, lateral and to some extent the dorsal wallof the laryngeal sack (" B-group "). These fibres seem to have no insertion
to any cartilage, terminating within the muscular wall itself. The ventral
portion of them, running caudally and mediad, contrary to the caudal andlateral direction of the A-group, is interwoven in the median line between
both sides~
Such a division of superficial and deep layers is however not so distinct, and they form altogether the very thick wall of the laryngeal sack.
For this reason, the muscles were taken as a whole by Carte-Macalisterand Benham as the thyreoarytenoid muscle, while Dubois regarded probably
the B-group as the lateral cricoarytenoid muscle.
It seems certain from my observatfon that the walls of the .laryngeal
sack are mainly formed by the thyreoarytenoid muscle, but this muscle
has, because of the extraordinary development of the sack, additional
insertions to the cricoid and tracheal cartilages. As to the problem, whether the B-group corresponds to the lateral cricoarytenoid muscle, I cansay nothing confidently. The bare fact is that the superficial fibres are
directed somewhat differently than the deep fibres.
In addition to A- and B-group, some muscle fibres situated m a morecranial place arise from the inner surface of the thyroid cartilage andcourse almost horizontally, surrounding the sack from lateral to reach the
middle ventral part of the arytenoid cartilage (" C-group "). We see besides
other fibres in a more. cranial level, connecting the basal part of the upper
process of the arytenoid cartilage with the root of the epoglottis (" Dgroup "). These groups are however not very clearly separated from eachother, but form as a whole the thick wall of the laryngeal sack. I guessthat the D-group in my description may correspond to M. ARYEPIGLOTTICus
mentioned by Dubois and Benham respectively in the larynx of B. Sibbaldii
and B. rostrata, or the superior and inferior arytenoepiglottic muscle
described by Carte and Macalister in B. rostrata, while the C-group seems
not to have been specially remarked by prevfous authors. They have pro-

33

On the Cetacean Larynx

bably ·considered this group merely as a part of the thyreoarytaenoideus.
The THYREOEPIGLOTTIC muscle, which extends between the root of the· epigl?ttis and the. cranial border of the thyroid cartilage, makes also a partof the muscular walls of the laryngeal sack.
Innervation of the laryngeal muscles (fig.18) :Of all the laryngeal muscles in the Sei whale only the cricothyreoideus
receives the external branch of the N. LARYNmcus CRANIALis. This nervec;lescends along the lateral surface ·of the larynx and, changing its course
slightly mediad, crosses over the thyroid cartilage. It comes down along
the lateral ridge of the cricothyreoideus, and then terminat.es there. . Theinternal · branch of this nerve runs through the thick laryngopharyngeal
muscle and ends in the mucous membrane of the larynx.
The N. LARYNGrnus CAUDAUS is considerably thick, ascends as the n ..
recurrens in the furrow between trachea and resophagus, comes near to
the inferior horn of . the thyroid cartilage and, crossing over the here
arising m. ceratocricoq.rytaenoitleus dorsalis, passes under this horn, to reachthe laryngeal sack, where it is divided into several branches, some of which
innervate the m .. cricoarytaenoideus and others enter .into the thick wallsof the sack, ending in its· muscles. The ending occurs in such a .way,that the uppermost branches course under the caudal border of the thyroid
cartilage and attain. the m. arYtaenoideus transversus; the next branches goto the C- and D-group ; the lowermost _branches, running in the caudal
direction, terminate in the superficial and deep layers of A- and B-group,
and we can trace their small branches macroscopically up to the half levelof ·the sack.
· Thus it is certain that, as all the muscles of the walls of the laryngeal
sack are innervated by n. laryngicus caudalis, A-group of muscles, in spiteof its insertion to the cricoid and tracheal cartilages, does not belong to
the cricothyreoideus.

.

Sperm whale :Extrinsic muscles
M. 1 STERNCYTHYREornEus and M. THYREOHYornEus: Connecting placesof these muscles to the thyroid cartilage are shown in t.he accompaning
figures (fig. 1'-a, b).

34

Hiroshi Hosokawa

M. LARYNGOPHARYNGICUS : Of this muscle the thyreopharyngeal' partis well developed, but the cricopharyngeal part is not found at all.
M. HYoEPIGLOTTJcus: This unpaired musde arises from the hyoid bone·(probably from its median part) and inserts to the ventral angle. of theepiglottic cartilage.
M. PALATOPHARYNGICUS : The aryteno-epiglottideal tube, protruding
into the choana, is encircled at the height of . its neck by a muscularbundle, which lies within the choanal walls. This bundle, named by vonBaer (1826) as the palatopharyngeal muscle, is well developed also in the
Sperm whale. It must have an important meaning in relation to the function of the aryteno-epiglottideal tube. For, when it contracts, the communication between the naso-pharynx and the laryngeal cavity is shut out ; atthe same time the aditus laryngis must be closed completely. The meaningof such anatomical structures will be considered later.
Intrinsic muscles( 1) M. CiircOTHYREOIDEUS: Stannius (1M8) and Rawitz described thismusole in phocaena communis as a small muscular bundle ; also Dubois
and Macalister (1867) remarked the same in Hyperoodon and Globicephalus
svineval respectively. But according to Benham, this muscle was invisible
in Cogia. In the Sperm whale, too, I could not ascertain its existence.
Whether it is quite absent in this whale, I want to determine by further
researches.
( 2 ) M. CRicoARYTAENomEus DoRSALrs : (fig. 6') ·This is very well developed.
As in the Sei whale not only the dorsal surface of the cricoid · cartilage
but also the inferior horn of the thyroid cartilage gives rise to this muscle(m. ceratocricoarytaenoideus dorsalis).
Its partial connection with theinferior horn was already reported by Stannius and Dubois, but Rawitz
stated nothing about it.
I was much interested by the asymmetrical development of thismuscle; namely it is very massive on the right side, but much poorer,
though dilated considerably, on the left side. It must have some intimate
relation with the asymmetry of the skull and nasal passage of this whale.
( 3 ) M. ·ARY'.rAENOIDEus TRANSVERsus: (figs. 6', 7') This muscle, connecting
transversely the arytenoid cartilages of both sides, is well developed. Its
lowermost fibres insert to the cranial border of the cricoid cartilage.

On the Cetacean Larynx

35

The caudal boundary of this muscle is distinct against the upper fibres ofm. cricoarytaenoideus dorsalis, the latter covering the former to someextent. This fact coincides well with Dubois' description on B. physalus.
As mentioned· already, the cricoidal insertion of m. arytaenoideus transversus
is also present in the Sei whale : This reJation, which ·according to· Fiirbringer (1875) never exists in the human larynx, seems to occur inthe whales not rarely.·( 4 ) M. CRICOARYTAENOIDEUS LATERALIS : This muscle was described by
Stannius in Phocaena, by Murie in. Globicephalus melas and by Benham
But Meckel .and Furbringer denied the existence of it inin Cogia.
Delphirtus delphis. .Watson and Young did not mention this muscle in
Beluga leucas (cited from Dub~is). Denying its existence in Hyperoodon
and in. other toothed whales, Dubois said that the so-called m. cricoarytaenoideus lateralis described by Stannius and Murie is nothing but a ceratal
portion of the cricoarytaenoideus dorsalis. In the Sperm whaie I too can
not find this muscle.
I want to mention here my finding of a: probably new muscle, whicharises from the INNER surface of the body of the cricoid cartilage· apd
inserts to the INNER surface of ·the arytenoid cartilage (fig. 9'); Though
its attachments to the cartilages are not very firm. compared with· othermuscles, it is certainly a special muscle clearly discernible from the· surroundings. Its nature. cannot be determined at present; but the fact itself
is interesting that such an unusual cricoarytenoid muscle exists, deserving
the name, " m. cricoarytaenoideus internus ". Moreover there are several
muscular fibres scattered on the inner surface of the thyroid. cartilage, andI wondered whether the muscle above mentioned would be nothing but acollection of these fibres.. But this assumption is not very probable, for theinternal cricoarytenoid muscle is a bundle so distinctly isolated.
( 5) M. TIIYREOARYTAEEOIDEUS: (figs. 7', 8') This is a massive muscularbundle, which .arises from aµ area along the caudal notch of the thyroid
cartilage and inser~s to. the ventral and lateral surface · of the ·mtiscular
process of the aryten0,id cartilage. As to k>cality and form, this muscle
is not so unusual as in .the Sei whale, for the laryngeal sack is absent
in this whale.
( 6 ) M. TIIYREOEPIGLOTT1cus and M. ARYEPIGLOTT1cus:
Though Rawitz
denied the existence of these muscles in Phocaena commurtis, Benham

36

Hiroshi Hosokawa

found them in Cogia. In the Sperm· whale I see no muscle fibres, which
may deserve the names. The upper portion of the m. thyreoarytaenoideus
comes in contact with the p~tiolus epiglottidis, but shows no direct connection to this. Absence of these muscles might be explained from the very
. firm ligamentous c6nnection between epiglottis and thyroid cartilage, where
almost .no mobility seems to remain.
III. The laryngeal cavity
Sei whale:
The larynx of the Sei whale does not exhibit a tubal structure as in
the toothed whales. As shown in fig. 10, the aditus laryngis of the Sei
whale differs in its general form not much from that of tli.e human being.
Namely, the tongue-shaped epiglottis projects from the ventral surface ofthe pharyngeal cavity, with its apex directed somewhat upwards (about 12
cm high). From its lateral ridge on each side a fold of mucous membrane(plica aryepiglottica) js stretched, encircling the laryngeal aperture, dorsallyand downwards to· the arytenoid cartilage. The cranial processes of thearytenoid cartilages are connected between both sides by a part of mucous
membrane ; the apex, though projecting upwards a little, is situated much
lower than that of the epiglottis..
The laryngeal cavity ·is in this whale a relatively narrow space surrounded by the paired arytenoid cartilages and the single crieoid cartilage
behind. The mucous membrane, covering the inner surface, is generallysmooth, but has a few fine grooves. This cavity passes downwards behindthe caudal processes of the arytenoid cartilages, which are also connected
between both sides, and continues further with the tracheal cavity.
The laryngeal sack (figs. 11, 13-17, 18):
As related before, the laryngeal cavity has in the Sei whale a large
sack-formed appendage, which is to be named "saccus laryngis ventralis ".
It is on the ventral wall of the larynx, elongated downwards throughthe whole length of the trachea, the ventral clefts of cricotracheal cartilages
offering a space just fitted for this sack. The constituents of the thickmuscular wall of this sack were already mentioned in the precedent chapter.
The cavity within the sack is elongated longitudinally and communicateswith the proper cavum laryngis through a slit along nearly the whole lengthOn the Cetacean Larynx

37of the arytenoid cartilage. The inner surface of the sack has much folds
and grooves; as shown in fig. 11: There are many longitudinal folds inthe upper part, while the lower parts show many reticulated grooves andthe meshes are especially fine on the dorsal wall. We see many granularprominences here on the mucous membrane. In the ventral parts the folds'and grooves are considerably rough. Judging from this structure, it seems
doubtless that this sack is liable to extension and contraction in the living
whale.
It is naturally important to know the function of this highly contractile laryngeal sack. I will consider this problem at first historically.
The honour of the discovery of the laryngeal sack is ascribed to J.
Hunter, as this great anatomist stated in his study· on Balaenoptera rostrata, that a sack is present on the ventral surface of the larynx.
Since then the 'laryngeal sack has been observed by Eschricht andReinhardt (1866) in Balaena mysticetus and by Carte and Macalister, andBenham in Balaenoptera rostrata. Beauregard and Boulart (1882), who
examined the laryngeal sack in Balaenoptera musculus, B. physalus andBalaena antipodum, noticed · the difference that the sack in question is far
smaller in Bqlaenidae than in Balaenopteridae.
In the Sei whale Schulte(1916) touched this sack briefly.
The. fact that this laryngeal sack is present only in the whalebone
whales, was indicated for the first time by Eschricht and Reinhardt.
Dubois, comparing between B. physalus and several kinds of toothed whales,· ascertained this. Rawitz, too, mentioned this fact, when he studied thelarynx of Phocaena communis.
In: the meantime the problems, what is the anatomical nature of thelaryngeal sack and why this. sack exists only in the whalebone whales,
have been very differently answered. Dubois regarded this sack. ashomologous with the Morgagni' s ventricle. He came to this opinion byI
comparing the laryngeal structure of toothed whales, because these animals
have no laryngeal sack on the ventral wall of the larynx, but have a recess
on each side of the median longitudinal membranous fold ; and he explained
this recess as corresponding to the Morgagni's vent~icle. Dubois thoughtthat the laryngeal sack of whalebone whales is formed by the further
enlargement of the recess, extension of its walls and by development of
the thyreoarytenoid muscle.

Hiroshi Hosokawa

38'

But Benham doubted the homogeneity of the laryngeal sack of whalebone whales with the laryngeal recess of toothed whales, though he accepted
the correspondence of the recess with ·the Morgagni's ventricle. Such aproblem is very difficult to decide, for the two structures, both of which
are appendages of the laryngeal cavity, differ very much from each otherespecially in their relation to muscles and cartilages. On the other hand,
there are undeniable communities between them, in spite of the differencesin size, form and structure of the walls.
A key for settling the problem may probably lie in the fact, that thelaryngeal sack for Balaenidae and Megapteridae is not so large as that of
Balaenopteridae, showing, so to speak, an intermediate form between thelaryngeal recess and the laryngeal sack. · I myself ascertained the· smallnessof the laryngeal sack in a f etus of Megaptera nodosa.
Now, on the function of the laryngeal s3.ck there seem to be threepossibilities as follows.
First, it may be possible that this is an apparatus for preventing theentrance of water and food into the respiratory canal, as the contraction ofmassive muscles of this sack will make the laryngeal sack, at the sametime the larynx itself firm ?nd solid, so as to avoid mis-swallowing of alarge quantity of food with water into the larynx and trachea. Rawitz, forinstance, stated this opinion. Prof. Ogawa has also an opinion near but alittle different to this, for he thinks that the blow of air produced by contraction of the laryngeal sack will prevent the entrance of water and foodinto the respiratory canal.
The second possibility is, that this sack may concern with complete
utilization of oxygen in the inspired air. Schulte said in his paper on a
fetus of the Sei whale, " that by its contraction and relaxation. during
submergence, a circulation of air in the wide trachea and bronchi might
be set up, which would favor the absorption of oxygen by bringing the air'in these passages more rapidly into contact with the respiratory membranethan could be done by the usual diffusion currents ". Another very interesting opinion related to the second possibility was stated by Negus in his
book titled ;, The Mechanism of the Larynx'', that all the air-sacs communi~cated with the respiratory canal of animals must be for the " rebreathingof air". To explain the "rebreathing of air'', he took up the frog as an
example, saying " a frog kept under water for a prolonged period expels airJOn the Cetacean Larynx

39

from its lungs into its mouth and pharynx ; the used-up air which has beenin contact with the pulmonary epithelium mixes with the relatively unused
air which has lain in· the centre of the sac-like lungs: This mixed air,when blown back into the lungs, gives up a fresh supply of oxygen andtakes up CO,, so that respiration can be continued for a time without the
intake of· a fresh supply of air ". According to him, the laryngeal air sacs
not only of the aquatic· animals including sea-lion, whales etc., but also of
the terrestrial animals are for this purpose.
The third possibility is the relation of this sack with the phonation.
The Sei whale has no vocal cord in the larynx; its absence is also provedin other whalebone and toothed whales.
And so, it is generally believedthat the whales produce no voice. Some !iUthors however insisted upon
voices of whales (Schneider, 1795; Murie; Rawitz). Turner (1872) explained
the mechanism of phonation in Balaenidae as such, that the elongated caudal, processes of the arytenoid cartilages are drawn near to each other andvibrate by the strong exspiration of whales. This opinion was accepted byWeber and Dubois, and was, though denied by Rawitz, adopted again byGoppert in .f!olk's "Handbuch der vergleichenden Anatomie" (1937). If
such a mechanism of phonation be true, we must hold in mind the utilityof the laryngeal sack in this function.
At pre~ent it is impossible to determine, which of these tqree be thefact. Moreover the truth may lie outside of them.
· Relation· of the laryngeal sack with ·th.e internal structure of the brain(fig..19):
While the Sei whale has a large laryngeal sack, the Sperm whale has
not such a sack. Probably relating to this difference, a remarkable different
structure was found in the medulla oblongata of whales. Namely, thenucleus ambiguus is poorly developed in the pigmy Sperm whale, Cogia
breviceps.(Fig. 12'), while the same nucleus shows in the Sei whale such a
striking development that the name ''a,mbiguus " is very unsuitable for thisanimal. Prof. Ogawa (1948) has directed his attention for the first time tothis fact and ascribed the· e:xtraordinary ·development of this nucleus in the Sei
whale to the large laryngeal sack. For the massive muscular sheets of thissack are innervated by the vagus and it is generally believed that thenerves innervating laryngeal muscles arise from.the nucleus ambiguus.

40

Hiroshi Hosokawa

To make this problem more definite, we must examine in the future
the brains of Megapteridae and Balaenidae, for the laryngeal sack of themis, as already mentioned, much smaller than that of Balaenopteridae, butlarger than the laryngeal recess of toothed whales.
Sperm whale :· The entrance to the larynx is in this whale a transverse fissure at the
top of the aryteno-epiglottideal tube, which protrudes into the choana andis encircled at its neck by circular fibres of the palatopharyngeal muscle.
The larynx of the Sperm whale shows therefore the well known characteristic of various toothed whales (figs. 7', 10').
Figure 11' shows the inner surface of the· laryngeal cavity. In thissketch the whole· larynx except the epiglottic cartilage is cut in the medianplane and the right half is removed. The cavity communicates above withnasal. cavity through the laryngeal aperture and below · with the tracheal
canal, while the greater part of this cavity belongs to the aryteno-epiglottideal. tube. Along the median line of the ventral wall, we see· a longitudinalelevation covering the median crest of the epiglottic cartilage. The mucous
membrane of the laryngeal cavity has numerous fine furrows and folds.
Very fine furrows run longitudinally in the upper half of this cavity andbetween them many minute points or holes are seen in rows. The mucous
membrane,· which covers the medial surface of the arytenoid ·cartilage isquite smooth, lacking of furrows. In the lower parts, near the trachea,
there are fine reticular furrows and folds, while the part covering themedian crest of the arytenoid cartilage has oblique or looped furrows.
From the caudal end of the median crest a membranous fold (plica
mediana) runs to the cranial and ventral border of the cricoid cartilage
and we see here a triangular recess on each side of this fold. The furrows
are reltively rough in this part and many lacunae are present, which seemto .be aperture? of the lymphatic nodules.*This triangular recess was already refered to in relation to the laryngeal sack. The opinion that this recess would correspond to Morgagni'sventricle in other mammals was stated for the first time by Murie in his* The inner surface of the laryngeal sack of the Sei whale does not show such lacunae, but
I found many relatively larger lacunae on the ventral surface of the membrane connecting theupper ends of the qanial processes of the arytenGid cartilages of both sides. (fig. 12)

On the Cetacean Larynx

41

paper on Globicephalus melas. Watson and Young published the same
opinion for Beluga leucas (cited from Dubois) and Dubois also accepted it .
. Physiological meaning of the aryteno-epiglottideal tube : .
The opinion has been prevalent, that the toothed whales can swallow
and respirate at the same time owing to this peculiar structure, for .thealimentary and the respiratory canal can be in this way completely separated from each other. This opinion seems to have its origin in Camper(1820), and was supported by Milne-Edward (1860), Gegenbaur (1891),
Zuckerkandl (1898) (cited from Boenninghaus) and Negus (1928). But it
had not so definite basis and was opposed to by Boenninghaus fromanatomical and physiological reasons. According to him, all the mammals
must dilate its pharynx at deglutition and simultaneously the larynx must
be shut off as a reflex by contraction of the laryngeal muscles. My observations upon the larynx of the Sperm whale justify more the opinion of
Boenninghaus. I want to suggest, that, when the palatopharyngeal muscle
is contracted at deglutition to avert the entrance of food and. water into
the choana, the laryngeal aperture also must be closed, so there would
occur no respiratory action. Besides, I. suppose, the Sperm whale would
have no need to be able to swallow and respirate at the same time, because
this whale used to take its food in the depth of sea. Taking in mind thehigh pressure of water there, it seems very probable that such a tubal
apparatus is exclusively to avoid the mis-swallowing of food and water into
choana and larynx, rather than to swallow and respirate simultaneously~
Another somewhat different .opinion was offered by Huber (1934), whostated in his paper on the nasal passage of Tursiops, that the palatopharyngeal muscle around the aryteno-epiglottideal tube, together with. them. maxillonasolabialis around the blow-hole and the· muscular plug in theosseous nasal passage, . belongs to the blow-hole mechanism, which performs
the task to shut the nasal passage closely, so as to avoid the entrance ofwater into lungs, when the whale submerges into the depth of sea. At
any rate, it. is quite reasonable, also from the stan'dpoint of dynamics, toclose the laryngeal aperture by contractlng the neck of this tube by thepalato-pharyngeal muscle.
As an appendix to this work I wish to say briefly about the Phonation

42

Hiroshi Hosokawa

of Whales.
Neither the Sperm whale nor the Sei whale has the vocal cord in thelarynx. Absence of the vocal cord in whales has been written by many
authors, for· instance. by Meckel for Delphinus and Phocaena, by Cuvier,
Stannius, Milne-Edward etc. for various kinds of whales, and Hunter insisted
that the whales do not possess the ability of crying. But Schneider stated,
in opposite to Hunter, that he heard from a "Gronlandfahrer " about cryingof whales and· Murie said in his paper on Grampus rissoanus, " that somespecies of whales utter sound is now unquestioned, notably the species
under consideration and the ca'ing whale '(Globicephalus melas) ".
How can the whale then make sound without the vocal cord ? Murie
thought, taking the membranous folds above mentioned- two folds are said
to exist in the larynx of Grampus- as the substitute for the vocai cord,that the forcible blow or exspiration of the whale would vibrate these folds,
producing sound. He added furthermore that in some animals, for instance,
Hyomoschus . aquaticus and ·Saiga tartaria, the vocal cords are placednearly vertically on the sides of the larynx, similarly as in the cetacean
larynx.
Concerning the whalebone wha!es, Milne~Edward (1876) said that theywould be dumb, but· Turner, Watson, 'Young and Dubois accepted the· possibility of them to make sound. They explained that the caudal processof the arytenoid cartilage would be vibrated by the exblows of the whale,
producing thus voices. Dubois added furthermore that perhaps a similar
modus of phona:tion is possible also for Otaria gillespii. Though Rawitz mentioned, opposing to Turner etc., that it is impossible for a cartilaginous
substance to vibrate and produce voice, yet he believed the ability of
whales to make voice, especially of Megaptera boops. According to him,
the organ of phonation is in the whales not the arytenoid cartilage but
membranous folds formed temporarily in the . larynx or in the soft palate,especially during ·the rutting season.
So far I have rev:iewed the opinions and possibilities published byvarious authors on the phon·ation of whales, but, regret to say, that the
final s'3lution of this problem is far b3y.::n1 us. But it s~~ms doubtlessthat some dolphins can make sound, ·though it might be faint. Of thewhalebone whales I have not b~en told by any whaler that he heard thevoice of whales. During my Antarctic whaling trip (1917-1948), I had noOn the Cetacean Larynx

43

opportunity to experience the voice of whales. Even, when· they were
hurted by the harpoon, they did not cry at all. If they could produce
voice, I supposed, judging from the situation, that they would make voices
in such cases. Of course it is not impossible that the whales produce
sound and communicate between them especially during the rutting season.
"'
Even at that time the sound might be so faint that it does not deserve ·tobe called "voice " ; but it would be sufficient for the commun!ction
between them ; the vibration will be propagated in the water and received
by their sensitive· auditory organs. If so, there is no need for us to seek
for any special vocal organ in the larynx, for such a vibration could bep;roduced without the vocal cord from other parts of the body. At · any
rate it would be necessary to know the nature of the voice of whales, its
relation to the breeding season and so forth.
IV. Comparison of thelarynx between the.Sei and the Sperm whale
From the foregoing description I will point out the remarkable diff erences in the larynx of the Sei' and th<: Sperm whale.
-·

----

Thyroid cartilage·~---

Sei whale

Sperm whale

Relatively small, without the ·median angle.

Of a large size, with a remarkable. median angle.
·-

Epiglottic cartilage

Arytenoid cartilages

Well developed and elongated'upwards. 'Relatively small.
Caudal process is well developed· and its apex is connectedwith that of the. other side.

Cranial process is continuous
between both sides at the upper
end. The caudal process isabsent.

Cr\coid cartilagea~d

Ventral half is widely lacking
continuous downwards tothe tracheal cartilages.

Completely ring-formed.

Tracheal cartilages

Ventrallacking.

Completely ring-shaped.

parts

are

widely·-·-·Thyreoarytenoid muscle

Long ana. wide, forming thewalls of the laryngeal sack.

An ordinary muscle bundle.

Laryngeal sackOf very large size.

Lacking, but provided: with asmall recess on each side. ·Aryteno·epiglottideal tube

Lacking perfectly.

Well developed, typical forthe toothed whales.

Asymmetry of thelaryngeal structure

Almost symmetrical.

Somewhat asymmefrical (fo~example, development of m.
cricoarytaenoideus dorsalis); ·44Hiroshi .Hoso.kawa

From this table, it m,ay be concluded that the most important difference
lies in the remarkable development of the laryngeal sack of the Sei whale
and in the. typical aryteno-epiglottideal tube in the Sperm whale; For,almost all of other differen~es are mor~ or less in relation to these features,
which are at the same time·the characteristic respectively of the whaleboneand of the toothed whales.
As the last problem, I wish to consider, why such a tubal structure of
the larynx exists only in· the toothed whales? And why is the laryngeal
sack present only in the whalebone whales? These problems, I suppose,cannot be answered merely froin the functional standpoint, but areconcerned in a high degree also with the phylogenetic relation. In otherwords such remarkable differences in the lq.rynx between toothed andwhalebone whales is not to be explained solely as· acquired characteristics
during the aquatic life.
The interrelationship. between these two groups of whales does not seemto be so close, and the resemblance of their external form might be rather
explained as a convergence of organisms ; . for, the inner structures of themare, in some points, too much different to be regarded as intimately related.
Concerning_the relationship between the toothed and the whalebone whales,
Dubois remarked that " der Larynx der Odontoceti steht dem urspriinglichen Typen naher, wahrend derjenige der Mystacoceti sich weiter
entfemt hat". Rawitz opposed to this opi,nion and said, "<lass Odontound Mystacoceten-larynx sich nicht aus einander entwickelt haben konnen,<lass der erstere nicht die Vorstufe des letzteren darstellt, <lass aber
Und daraus folgt, <lass sie
das umgekehrte Verhaltnis nicht statthat.
aber an der Wurzel nicht einander zusammenhangen, nicht direkter Verwandtschaft mit einander stehen ", and I myself agree with Rawitz's
opinion. Putter (1902) also stated approximate ideas in his comparative
anatomical work upon the eyes of marine m?-mmals.
Summary and Conclusions
1) As to the laryngeal cartilages and muscles. many remarkable differences.. are found between the Sei and the Sperm whale.
2) The· larynx of the Sei whale shows the mo3t noteworthy characteristic
in the presence of an enormous muscular sack on its ventral side, saccus
laryngis ventralis, and almost all of the peculiarities of its cartilages andOn the Cetacean Larynx

45

muscles are more or less concerned with this laryngeal sack.
3) Four muscle-groups (A, B, C, D) were classified in the muscular wallsof the laryngeal Stick, with considerations upon their homogeneity withthe laryngeal muscles of other mammals.
4) These muscle-groups of the laryngeal sack are all innervated by thecaudal laryngeal nerve.
5) The extraordinarily well developed nucleus ambiguus in the medulla
oblongata of the Sei whale. seems to be intimately related with the actionof this laryngeal sack, which is probably able to contract and dilate ina considerable degree. The corresponding nucleus is not well .developedin the toothed whales, which lack of ·the laryngeal sack.
6) The general characteristic of the larynx of the toothed whales, including
the Sperm whale, lies in the upward elongated epiglottis and arytenoid
cartilage, which form altogether the aryteno-epiglottideal tube.
7) Som.e asymmetrical relations were noticed in the larynx of the Spermwhale, for instance in the development of the dorsal cricoarytenoid muscle
and in the relative position of oesophagus to , the· larynx. These asymmetries niay be related with the extraordinarily asymmetrical nasal passages of this whale.
8) With the completely ring-formed cricoid and tracheal . cartilages, theSperm whale is, together with Cogia and· Tursiops, an exceptional casein the realm of Odontoceti, for these cartilages have a median cleft inthe ventral portion in almost all of the toothed whales.
9) The lateral cricoarytenoid m.uscle was not found at all in the Spermwhale, but I saw a muscle bundle, probably not yet known, deserving
the name " m. cricoarytaenoideus intern us ".
10) The remarkable differences in the larynx between the 'whalebone andtoothed whales cannot be e:xplained merely from the adaptation of thesewhale to the aquatic life, but must be rather refered to the phylogenetic
relation ; namely, I believe, coincidently with some previous authors, thatthe two groups of whales, whaebone and ·toothed whales, are not so
intimately kindred to each other as genera1ly assumed.

Hiroshi Hosokawa

46

Explanation of figures :Sei whale
Fig. 1,Thyroid cartilage

Fig. 2,

Epiglottic cartilage

Fig. 3,

Cricoid and tracheal cartilages

Fig. 4,

Arytenoid cartilage

Fig. 5;

Larynx (ventral view)Fig. 6,

Larynx (dorsal view)Fig. 7,

Larynx. (dorsolateral view)Fig. 8,Larynx (the ventral ~ight quarter is removed)

Fig. 9,Median section through the larynx

Fig. 10,

Laryngeal aperture

Fig. 11,

Laryngeal sack (inner surface)Fig. 12,

Lymphatic lacunae on the ventral surface of the membrane connecting the cranial

Fig. 13,

Horizontal section through the larynx (1)Fig. 14,

Horizontal section through the larynx (2)processes of the arytenoid c.artilages

Fig. 15,

Horizontal section through the larynx (3)Fig. 16,

Horizontal section through the larynx (4)Fig. 17,

Horizontal section through the larynx (5)Fig. 18,

Innervation of the laryngeal muscles

Fig. 19,

Nucleus ambiguus

Sperm whale. Fig. 1'
Fig.

'

21;

Thyroid cartilage
Epiglottic cartilage

Fig. 3',Fig. 41

Cricoid cartilage

Fig. 5',

Tracheal cartilages

Fig. 6',
Fig. 7',

Larynx (dorsal view)'

Arytenoid cartilage

Larynx (dorsolateral view).

Fig. 8',Fig. 9',M. cricoarytaenoideus internus

Fig. 101,
Fig. 11',

Laryngeal aperture
Laryngeal cavity

Fig. 121,

Nucleus ambiguus of the pigmy Sperm whale (Cogia breviceps)

Larynx (lateral view, the right lamina of the thyroid cartilage is removed)On tp.e Cetacean Larynx

47· Literature. cited· l) . Tyson,E.;

Phocaena, or the anatpmy of a porpess, dis!Jected at Gresham eollege etc. London.

1680. (cited from Rawit~)2) Hunter, J.; Observations on the structure and oeconomy of whale~. Phi!os. Transact. Roy.
Soc. London. Vol. 16 (1785-90). 1787. pp. 306-351.
3)

Schneider, J. G.; Beitrage zur Naturgeschichte der Walfischarten. Leipzig. 1795.

4) Camper, P.; Observations anatomiques sur la structure interieure et le squelette de plusieurs
especes des Cetaces. Paris. 1820. (cited from Rawitz)
5) Baer, K. E.; Ueber den Braunfisch (Delphinus phoq1ena). Isis von Oken. Jahrgang 1S26. Bd.
1. Heft 1-6. 1826. (cited from Rawitz)
6) Meckel, J. F.; System der vergleichenden Anatomie. 6. Teil. 1833.
7) Cuvier, G.; L~o~s d'anatomie co:n;lparee. Tome 4. Premiere partie. 1835.
8) Stannius, H.; Lehrbuch der vergleichenden Anatomie. 2. Tei!. 1846.
9) Stannius, H.; Beschreibung der Muskeln des Tlimmlers (Delphinus phocaena). Muller's Archiv
flir Anatomie 1849. (cited from Dubois)
10) Eschricht, D. F. and Reinhardt,].; On the Greenland Right-whale (Balaena mysticetus). Ray
Society. Recent memoirs on the cetacea. 1866. pp. 1-150.
11) Macalister, A~; On some points in the anatomy of Globicephalus svineval (Gray).

Proc.

ZooL Soc. London. 1867. pp. 477-.-482.
12) Burmeister.; Preliminary observations on the anatomy of Pontoporia blainvillii. Proc. Zool.
·Soc. London. 1867. pp. 484-489.
13) Burmeister ; Preliminary Description of a new species . of Finner Whale (Balaenoptera
bonaerensis). Proc. Zoo!. Soc. London. 1867. pp. 707-713.
14) Turner,W.; A contribution to the anatomy of the Pilot Whale (Globicephalus svineval,
Lacepede). Journ. Anat. Physiol. Vol. 2 (second series, Vol. 1). 1868. pp. 66-79.
15) Carte, A. and Macalister, A.; On the anatomy of Balaenoptera rostrata; Philos. Transact.
Roy. Soc. London. Vol. 158. 1868. pp. 201-261.
16) Murie, J.; On Risso's Gramus, G. rissoanus (Desm.). Journ. Anat. Phrsiol. Vol. 5. 1871. pp.
118-138.
17) Turner,W.; An Account of the Great Finner Whale (Balaenoptera Sibbaldii) stranded atLongneddery. Part 1. The soft parts. Transact. Roy. Soc. Edinburgh. Vol. 26. 1872. p.197.
(cited from Dubois)18) Murie, ].; On the Organisation of the Chaing whale (Globicephalus melas). Transact. Zoo!.
Soc. London. 1874. Vol. 8. p. 236. (cited from Dubois)19) Milne-Edwards, H.; Les:ons sur la physiologie et l'anatomie comparee de l'homme et desanimaux. Tome 2. Paris. 1876.
20) Flirbringer, M.; Beitrage zur Kenntnis der Kehlkopfmuskulatur. Jena. 1879.
21) Sanctis, L.; Monografia zootomico zoologica sul capidoglio arenatci a Porto St. Giorgio. Atti
de Lincei Mem. Cl. Sc. fis. ecc. Ser. 3. Vol. 9. 1879. p. 160. (cited from Dubois)
22) Watson, M. and Young, A. H.; The anatomy of the Northern Beluga (Beluga catodon, Gray,s. Delphinapterus leucas Pallas) compared with that of other Whales. Transact. Roy. Soc.
Edinburgh. Vol. 29. Part 1. 1878-79. (cited from Dubois)

Hiroshi Hosokawa

4823) Ho'!'les, G. B.; On some p9ints in the anatomy of the Porpoise (Phocaena communis). Journ.

Anat. Physiol. Vol. 14. 18'/9. pp. 467---:-474.
24) Beauregard, H. et Boulart, R.; Recherches sur le larynx et la trachee des Balenides. Journ.
de l'anatomie et de la physiologie. Vol. 18. 1882. pp. 611-634.
25) Weber, M.; Studien iiber Sjiugethiere, ein Beitrag zur Frage nach dem Ursprung der Ceta-

ceen. Jena. 1886.
26) Dubois, E.; Ueber den Larynx (der Cetaceen). Weber's "Studien iiber Saugethiere ". 1886.

pp. 88-111.
27) Rawitz, B.; Die Anatomie des Kehlkopfes und der Nase von Phocaena communis Cuv.

Internat. Monatsschrift fiir Anat. u. Physiol. Bd; 17. 1900. pp. 245-354.
28) Benham, W. B.; On the larynx of certain whales (Cogia, Balaenoptera and Ziphius). Proc.

Zoo!. Soc. London. 1901. I.. pp. 286-·300.
29) Boenninghaus, G.;

Der Rachen von Phocaena communis, Less. Zoo!. Jahrbiicher. Bd. 17.

1902 (1903). pp. 1-98.
30) Putter, A.; Die Augen der Wassersaugethiere. Zoo!. Jahrb. Bd. 17. 1902 (1903). pp. 99-392.
31) Schulte, H. w. von; Anatomy of a fetus of• Balaenoptera borealis. Memoirs of the American

Museum of National History. New Series. Vol.. 1. Part 6. New York. 1916. pp. 389-502.
32) Negus,; The mechanism of the larynx. London and Tombridge. 1928.
33) Huber, E.;

Anatomical notes on Pinnipedia and Cetacea. Contributions to Palaeontology
IV. Carnegie Institution of Was~ington, Publica~ions. No. 447~ 1934. pp. 106---136.

34) Bolk und Lubosch; Handbuch der vergleichenden Anatomie der Wirbeltiere. Bd. 3. Kehlkopf.

(by Goppert). 1937.

i

49

On the Cetacean Larynx

Fig. 1.

Cart. thyreoides (1/8)

ventral viewM. thyreohyoideus~

dorsal view

Incisura thyreoidea cranialis
Cornu
hyoideumM. thyreopharyngicus

A

Incisura thyreoidea caud.

cCornu cricoideum ---Fig. 2. Cartilago epiglottidis (1/4)
dorsal viewlateral view

~ '\

I ..

!)
~
Fig. 3.

Cart. cricoides et Cartt. tracheales(1/8) dorso-lateral viewM. cricoaryt; dorsalis- - Facies artic. arytaenoidea//

A (M. thyreoaryt.)
Fades artic. thyreoidea

Petiolus epiglottidis

M. cricothyreoideus

Proc. lateralis
M. cricopharyngicus

Cartt. tracheales

50

Hiroshi Hosokawa

Fig. 4.

Cart. arytaenoides (dextra)(1/8)

dorsal view

ventro-lateral view

medial viewProc. cranialis
M. arytaenoid.
transversusD(M. aryepiglotticus)·M. cricoaryt.. "dorsalisc·,--·-··- M. cricoaryt.
'\l'll"'ll·.-..._
dorsalis(M. vocalis)A(M. thyreoarytaenoid.)

Fades articularis cricoidea

B(M. cricoaryt.
!at.)Proc. caudalis

Fig. 5.

Larynxventral view

(1/6)

M. thyreohyoideus.. - M. sternothyreoideus___ M. thyreopharyngicus

Fig. 17

Fig. 16

M. cricoaryt. dors. _-Fig. 14\ __ - - -· M. cricothyreoideus- - -A/+----Fig. 13/

Fig. 13---;,/(M. cricoaryt. lateralis)(M. thyreoarytaenoideus)

Trachea

Saccus laryngis

On the Cetacean Larynx

Fig. 6.

Larynxdorsal view (2/15)

Raphe pharyngis

Oesophagus

Fig. 7.

LarynxOesophagus,M. cricopharyngicus

M. cricopharyngicus

dorso-lateral view (2/15) -51

Hiroshi Hosokawa

52
Fig. 8.

Larynx(the ventral right quarter is removed) (2/15)

Plica aryepiglottica

Cart. epiglottidis (Petiolus)

Cart. thyreoides

C (M. vocalis)

TracheaFig. 9.

Saccus laryngis

Median section through the larynx (1/8'
Cart. epiglottidis

Cart. thyreoides

c

'

I

'

'

I

Oesophagus Trachea Saccus laryngis.
· On the Cetacean Larynx

Fig. 10,53

Aditus laryngis (1/6)

Fig. 11. Saccus laryngis
inner surface (1/6)- - Cavum laryrigis

Fig. 12.
Lymphatic lacunae on the ventral
surface of the membrane connecting the
cranial processes of the arytaenoid carttiages (1/4)

Hiroshi Hosokawa

54

Horizontal section through the Iarynx.
(1) (2/9)

Fig. 13.

~esophagus

A (M. thyreoarytaenoideus)Fig. 14.

C Horizontal section t hrough the larynx (2)
avum
Oe h
(2/9)
~~s a· M. cricopha·
Cart. arytaenoides ' ' laryngis
ryngicusM·\. cncoarytaenoid. dors
I.

- Cornu cricoideum(Cart. thyreoid.)

B
I'

Saccus 1a'ryngisAOn the Cetacean Larynx

Fig. 15.
Cavum laryngis

55

Horizontal section through the larynx (3) (2/9)
Oesophagus

M. cricothyreoideus\\

Cart. thyreoides

Fig. 16. .
Cavum laryngis

Saccus laryngis

Horizontal section through the larynx (4) (2/9)

Oesophagus

M. arytaenoideus
transversus Cart. arytaenoides
/

/

/

M.

thyreopharyngicus"".l::::,..l'--:~,

S~ccus laryngis

56

Hiroshi Hosokawa
Horizontal section through the larynx (5) (2/9)

Fig. 17.

\\'

Plica aryepiglottica

Cart. thyreoidesD- (M. aryepiglotticus)

Innervation of the laryngeal muscles (2/15)
Fig. 18.
N. vagus-N. laryngicus cranialis (Ramus extenus)

M. thyreopharyngicus -·M. cricopharyngicus- -M, ceratocricoaryt. --- ·K
dorsalis
.·•

,.,
/

M. cricothyreoideus•/' ' Saccus laryngis
N. laryngicus caudalis

On the Cetacean Larynx

Fig. 19.

Medulla oblongata of the Sei-whale (5/2)

X

Fasciculus solitarius'

57

Fasciculus longi- Nd. n.
Nd. dorsalis
tudinalis medialis hypoglossi
n. vagi

:xn

\

lX

Corpusrestiforme

-

'\

/'/

Radix n.--/glossopharyngici
1
Radix n. vagi

I

1

I

Nd. ambiguus\ Tractus, spinalis
n. trigemini/

Fasciculi
Nd. olivae Nd.
Ne!. Nd. tract. spinalis
pyramidici accessorius olivae ambiguus n. trigemini
medialis
Fig. l'b Cartilage thyreoides (Lamina dextra) (1/8)
outer surface

Fig. l 1aM. thyreohyoideus- .... - ... Foramen

Cartilago thyreoides (Physetercatodon) seen from the ventroIateral side (1/8)
Incisura ~.thyreoidea cranialis

thyreoideum,
M. sterno- i!
thyreoideus

inner surface.

M. thyreopharyngicus
J

Epiglottis, -......

Incisura thyreoidea caudalis
M. thyreoarytaenoide us' '- - - - -M. ceratocricoarytaenoid.
dorsalisHiroshi Hosokawa

58

Fig. 21a Cartilago epiglottidis (1/6)
ventro-lateral view · dorso-lateral viewFig. 21b Cartilago epiglottidis (I/6)
lateral view .
horizontal section

Syndesmosis withthe thyroid cartilage
Fig. 31
dorso-lateral view

Cart. cricoides (I/G)
ventro-lateral viewM. cricoarytaenoideus internus

Proc. cranialis

Proc. cranialis - - - - - - -M. arytaenoideus transversus - - - - - - -.....

M. cricoaryt. dorsalis - - - - -,f;.L-.,
M. crycoaryt. internus ---- -/r - - M: thyreoarytaenoideus'

Facies artic. cricoidea

59

On the ·Cetacean Larynx

Fig. 4'bCart. arytaenoides .(sinistra) (2) (1/6)

lateral view

medial viewFig. 51a. Cartilagines
tracheales (1) (1/6)
ventro-lateral view

' - - - - - M. arytaenoideus transv. - - - -\
I
I

t'

',

·-- -M. cricoaryt. dorsalis

dorsolateral view

-J

Facies artic. cricoidea
M. crycoaryt. internus - -Cart. trachealis I'---M. thyreoarytaenoideus

Fig. 5'b Carttilagines tracheales (2) (1/6)
median section
Fig. 61 Larynx (2/9)
ventral -dorsal viewM. thyreoary- Cart. ary- Cart. epi- M. arytaen.
taenoideus
taenoides
glottidis transversus
/

/

/
/

/

.
M. ceratocricoarytaen. dorsalis

Hiroshi Hosokawa

60

Fig. 7'

Larynx (2/9)
Aditus laryngis

Cartilago arytaenoides

Tuba aryepiglottica

Cartilago thyreoides- Cartilago
thyreoides(Lamina
dextra)- - - M. thyreoarytaenoideus

Cartilago
cricoides.

M. cerato- ___ ,,cricoarytaenoideus
dorsalis/,

Fig. 8'Larynx (1/6)- - .,. Aditus laryngis- - -- Tuba aryepiglottica

Cart. cricoides __ _Cartilago
thyreoides(Laminasinistra)

//
.c.-/

M. cricoaryt.
dorsalis
I

I

Fades articularis thyreoidea

M. thyreoarytaenoideus

On the Cetacean Larynx
Fig. 9'.61

M. cricoarytaenoideus internus (1/6)
medial view
~ventralCart. arytaenoides

M. cricoaryt. internus

Cart. cricoides - - - -Fig. 10'Aditus laryngis (1/6)_____ Cavum nasi--- - - Aditus laryngis------ Tuba aryepiglottica
M. palatopharyngicus- Cavum pharyngis

Oesophagus
M. cricoarytaenoid. dorsalis
TracheaHiroshi Hosokawa

62

Fig. 111

Cavum laryngis (1/6)+

dorsalcaudalrostralventralAditus laryngis

Cartilago arytaenoides(sinistra)
M. arytaenoid. transversus ,M. cricoaryt.

Cart. cricoides -Trachea-

Cart. cricoides /1
Recessus.
M. thyreoarytaenoideus Plic~
mediana laryngis
Fig. 121 .

Cart. thyreoides

Medulla oblongata of the pigmy Sprm-whale (10/3)
X

XIl

Nd. n. hypoglossi

Fasc. longit. medialis
I

I

/

/

....

Fasciculus ------~--,
solitarius-..... --·--·--·.Nd. dorsalis
n. vagi

Corpusrestiforme- -Tract. spinalis
n. trigemini
\
\

. .
./
Rd
a rx n. vagi

/

I

\

' '·----Nd. tractus
spinalis n.
trigeminiI

Ne\. ambiguus

Nd. oli~ae acces- Nd. oliv~e Nd. ambiguus
sorius medialis

Bacteriological Studies on Freshness of vVhale Meat
(Report No. 1)
Tomoichiro Akiba, Takeshi Tsuchiya,
Makoto Umehara and Yoshiharu Natsume
In using whale meat and organs as food-stuff and material of medicaments the value of their utilization depends to a large extent on theirfreshness. Consequently, it is of great importance to take necessary steps
preventing decrease in freshness, and in other words the factors causing
deterioration must be clear. However,· the research along this line has
been insufficient. In order to study these points it is necessary to investigate the various factors such as whether this is due only to self-digestion
or autolysis after death of whale, or besides what route bacteria take to be
involved in this, or what kind of bacteria invade when and how, if suchbacteria are involved in this. We began this research in the summer of1947 with this object, but our material was very scarce in 1947 due to thepassed fishing season,' and so these results of .1947 and a part of 1948 will
be consolidated and reported as Report No. 1.

Experimental Results
1. Collection of Samples. Trips were made twice, onoe in August, 1947
and another in July, 1948 to Ayukawa-machi which is located at the tlp of
Ojika Peninsular. The determination of various kinds of bacteria, putrefaction tests, etc. in this report were carried out on these samples takenfrom one whale which was dissected in August 1947. As materials for our
tests, its blood, muscles, contents of small and large intestines and sea
water were used. Blood which flowed out from the heart during dissection,
was collected with a sterilized pipette. Consequently, since dissection was
not carried on under sterilized condition, the invasion of bacteria from theenvironments cannot be absolutely ·refuted. However, since a sterilized
pipette was inserted deeply in a copious flow of blood, it may be assumed
that bacteria contained in the blood itself were collected. As to meat,
about 20 c. cm. of meat lump was cut off, placed in boiling water and boiled
for about 3 to .5 minutes. After sterilizing the surface of muscle tissue itI64Tomoichiro Akiba, Takeshi Tsuchiya, Makoto Umehara and Yoshiharu Natsume

was cut open with a scapel, and 2 or ,4 gm. of muscle tissue was cut off
from the center. To this, 8 or 2 cc. of physiological saline solution wasadded and stirred well. After standing only the meat juice was taken andused in the experiments. As to the contents of the intestines, these were
cut open and the contents were collected with a sterilized pipette. Thiswas diluted to five times with physiological saline solution and used in theexperiments. The intestines were also ·swollen due to gas produced byfermentation of the contents at that time. Sea water was collected in frontof the works and from the bay.
As culture media sea water agar media, meat infusion agar, Endo's
media and cysteine glucose agar were used. Sea water agar media wereused for the purpose of not losing specific sea water bacteria, cysteine agar
media for anaerobic bacteria, and Endo's media for the isolation of colon.type bacteria. Sea water agar media were made according .to Zo Bell's
method, and 0.5% of peptone, 0.01% of ferric phosphate and 1.5% of agaragar were disolved in 1000 cc. of sea water being aged before the use.
2. Freshness and Number of Bacteria. Number of bacteria detected inblood and muscle was almost proportional to freshness and the number ofbacteria also increased as the time elapsed longer after catching. However,Table 1.

Freshness and Number of Bacteria

Date Whale Caught
Time of elapse after catching (hours)

<Ll
OS

&l::I

'$
~

.c

g
::s

z

Sei
Whale

B 3
Sei
Whale

B 4
Sperm
Whale

Aug. 12·July 25

July 25

July 25

July 27

30

34

26

68

24

Fresh
(80%)

Fresh
(80%)

Satisfactory
(50%)

Fresh
(80%)

70,00012,0009,360,000

23,400

9,400

8,200

5.9
6.0

5.6
6.1

Blood

Meat influsion agar media

per ml.

Sea water agar media

Muscleper gmLarge Intestinecontents
per ml.
Small Intestinecontents
per ml.

Meat influsion-agar media
Sea water agar media
Meat influsion-agar meiiia

SeveralSea water agar media

0

Meat influsion-agar media

SeveralSea water agar media

Several

·;:::

t)

B 1
Sei
Whale

Satisfactory(50%)
Innumerable
Innumerable
4,575
4,425

Freshness

OS

A 1
Sperm
Whale

B

2

Blood
6.2
6.2
Muscle6.2
5.8
Note : Freshness was expressed by the visual judgement of experts.

pH

IBacteriological Studies on Freshness of Whale Meat65

it was not clearly recognized that there was some relation between freshness and hydrogen ion concentration in blood and muscle at the time ofdissection, but all the reactions indicate the progressing decomposition of
the sugar.
3. Indentification of Isolated Bacteria. In this paper are reported only
the results of experiments as to the samples collected from a sperm whale
dissected in August 1947. Result of another samples collected in July 1948
shall be reported in the second report. Identification of various bacteria
was attempted as to bacteria isolated at the works and also bacteria isolated after bringing- samples to our laboratory. Even the bactetia obtainedby anaerobic culture grow better by aerobic culture, after successive
transfer on culture media, so all the succeeding examinations of biologicalcharacters were made by aerobic culture.
Microscopic examinations were made as to the pure culture of the 57
strains to determine their shape, grouping, Gram's stain, size, motility, etc.
and furthermore 19 . species were selected according to shape, form andlcolor of colonies on the media such as meat infusion agar, sea water agar
media and cysteine glucose culture, and gas production, liquefaction ofgelatin, hydrogen sulphide production and indol reaction. As to 19 strains
the further examinations were made as following table (Table 2).
Of 19 strains another three were same strains, so that 16 species wereidentified according to Bergey's Manual of Determinative Bacteriology andother literatures. It was possible to determine 9 different Micrococci, 1
Streptococcus foecalis, 2 Flavobacteria and 4 Achromobacters.
These bacteria may generally be found in soil, air;· water, etc., andalthough there may be few cases in which their biological characteristics do
not exactly coincide with those found in references, but the principal ones
coincide exactly and . we could not find new species. In a case as MicrococClJS epimetheu~, there is a question more or less compared with the place
primarily found, but there is not any reason that such bacteria must not
be found in places other than where they were found, so that it may be
assumed that the presence of such bacteria is probable.
4. Putrefaction Tests. When these bacteria were inoculated on pieces ofwhale meat and left standing at 30°C, 8 strains produced rancid irritatingor light deterioration odor after 18 hours, and 14 strains after 48 hours, andthe same time pH trended to alkaline.

Tomoichiro Akiba, Takeshi Tsuchiya, Makoto Umehara and Yoshiharu Natsume66

Also, in case several bacteria being grouped according to the origines
where they are found, are inoculated on the surface of a piece of meat,
the rancid odor is produced faster than in the previous cases and appears
after 6-12 hours.

~

Before

19
88
15
F 17
F
5
6
F
1
F
F 65
F 66
F 12
c 32
c 33
c 34
B 24
B 82
I 43
I 44
I 45
s 90
s 97
Control

6.2
6.2
6.3
6.2
6.3
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
- '6.2
6.2
6.2

F
F
F

Note:

I

Table 3.

Deterioration Test, -pH aod Odor

After
4
hrs.

6

6.1
6.2
6.1
6.1
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.1
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2
6.2

6.0
6.0
6.0
6.3
6.2
6.0
6.1
5.9
6.0
6.0
6.1
6.3
6.0
6.0
6.0
+6.2
6.2
6.0
6.0
6.0
6.2

I

24

48

6.8
7.0
+5.9
6.4
+7.0
6.4
+6.8
6.4
6.3

6.9
7.4
it-6.66.5+7.4
it-6.8it-6.8±6.6
it-6.3
+6.4
±7.6
6.1
+6.2
+7.4
+7.4
tft7.8
5.8
5.8
±6.7

26

I

6.0
6.0
6.0
6.3
6.2
6.0
'6.1
5,9
6.0
6.0
6.1
6.3
6.0
6.0
6.0

6.3
6.4
+6.0
6.3
±6.4
6.0

it-6°3
6.2
6.0
6.0
6.0
6.2

it-6°6
6.0
6.0
6.0
6.0
6.2

I

±6.4
6.2
6.1
+6.0
6.1
6.1
+6.0

+6.4
6.5
6.0
+5.7
+6.8
7.0
it-6.9
5.8
6.2
5.6
6.3
6.3

±9.4
ti.6

±~.5

6.3

7,7
it-8.2it-8.2it-7.0tft8.0tft7.6it-7.2
+6.8
+6.4
+6.6
tft8.0it-7.0fu6.8
it-8.0it-7.7
tft8.06.6+6.0
I +7.4
I it-7.0
I
I 6.6+-tit indicate the degreee of deterioration odor.

~

Before

After
4
hrs.

F groupC group
B group
I groupS groupControl

6.2
6.2
6.2
6.2
6.2
6.2

6.2
6.2
6.2
6.2
6.2
6.2

B
rial gro ps

I

'

18

12

6

12

18

24

48

Oder

6.4
6.2
6.3

+6.4
6.4
+6.3
+6.3
6.2
6.2

+6.S
6.5
+6.4
+6.5
6.3

it-7.8tft6.2+6.8
fft8.2tft8.0+7.4
6.4

Putrefied odor
Putrefied odor
Ester-like odor
Putrefiedirritating odor
Putrefied odor

+6.6
6.2
6.2

6:2

±6.4

I

it-7.0
+6.8
+6.8
6.2

If 0.2 cc. of bacterial emulsion injected into the p:eces of meat, theproduction of odor is even more conspicuous and als9 pH becomes conspicuously alkaline.
5. Invading Route of Bacteria. The route of entry of bacteria into
.blood and muscle has not been clarified as yet, but 4 routes of their entry

Bacteriological Studies on Freshness of Whale Meat

can be assumed, that is, explosion wound .caused. by spear, compressed air
pumped into the whale after death in order to keep it afloat, respiratory
tract such as lung and the intestinal tract.
In the case of entry through the explosion wound, there is cons:derable
negative pressure in the severed vein on account of the bleeding, therefore
it is understandable that some bacteria sticking to the part affected will beabsorbed and reach to the heart. When pig is slaughtered and bacteriacoated on the wound (in such cases, it is obvious that easily detectablebacteria are used), it is reported that the same bacteria can be found in
the bone marrow, and so these facts support this theory. Whales being
speared are especially different from animals being slaughtered, because
the former try to escape with strenuous activity for more than one hour.
More opportunity may be given for bacterial invasion due to the laboured
breathing during that time. Next, in regards to pumping in high pressure
air, special efforts to prevent entry of bacteria are not taken, so it can beassumed that there is a possibility of bacterial invasion. Also the cutting
open the abdomen to cool can give the path of entry for bacteria. Thirdly,in the case of respiratory tract, bacteria can be discovered in the lung of
human being and slaughtered pig, in which even the scraps of food arealso occasionally found, so there are some scholars who say -contam:nationof blood in heart is caused by passing through veins from the lung. If
the detection of principally soil bacteria in lung of pig by Htilphers is taken
into consideration, this path of entry can be naturally considered. Even
with the :present experiments, Micrococcus halophilus, which is a specificsea water bacteria, was detected in the sea water and muscle ; also
bacteria detected in blood and muscle· belong chiefly to soil bacteria; andsince a long time has elapsed between the time of catching and thedissection, it is considered that the above three routes of entry play considerably important parts.
However, since whales are animals living in the sea, there isn't any
reason why the entry of bacteria into blood and muscle by way of intestinal
tract cannot be considered. Even in the case of drowned or poisoned
human beings or animals, it can be assumed that perhaps the large numberof bacteria which originarily live in the intestines, enter into the blood vessels through the mucus membrane of the intestine due to the bacterial
permeability of the intestines after death and to the extraordinary increase. 68

Tomoichiro Akiba, Takeshi Tsuchiya, Makoto Umehara and Yoshiharu Natsumeof bacteria. In the case of whale this mode of entry maY be accelerated
by the air forced into the abdomen, by which the bacteria invade into theblood, and then they will circulate throughout the body and extend from theend of blood vessels into the muscle. ·Furthermore, from long experience,the staleness of fishes is prevented by removal of the intestine as soon as
possible after death, so it may be assumed that intestinal route plays a. major role. Also, from our experiments, the facts that 5 kinds of bacteria,
i. e. Micr6coccus piltonensis, , M. epimetheus, Flavobacterium sewanense,
Achromobacter ubiquitum and Achromoba'cter halophilum were found in
the contents of the intestine, muscle and blood, and that Streptococcus
faecalis which is always found in the intestine was detected in muscles,
supports the theory of the intestinal tract route. We were not able todiscover Bacterium coli in samples of 1947. This may be due to the factthat Bacter:um coli cannot exist for long in the blood as whale is a warm
blooded animal similar to pig. However, this requires further study in the
future to settle this point.
6. Putrefaction and Bacteria. It was proven that the number of bacteria
will increase in blood and muscle as degree of freshness decreases. Next
is that three strains of the detected bacteria do not grow at temperatures
above 35°C. The majority has optimum temperatures from 27-30°C, so if
the temperature of the body of a whale from the time of catching to dissection is within this range, it will be the optimum condition for increase ofbacteria and their putrefaction will occur faster.
Furthermore, these bacteria grow even in a media of low salt content(1%), but growth will be better at about 3%. If the concentration isincreased to 15%, growth will be greatly reduced, only 4 strains will exist
at 20% and growth is checked at 25%. Therefore, putrefaction can beprevented by maintaining salt contents above 25%.
Of the 57 strains of bact.eria, 38 decomposed glucose and produced large
quantity of gas. The large quantity of foam observed on stale whale meat..is probably due to such .bacteria. If such bacteria are inoculated on apiece of whale meat and its pH is exam:ined, at first it is about 6.2 andwill become lower after 24 hours, but after 48 hours it will become 6.48.2 and will give off rancid, irritating or at times a mild putrified odor.
These phenomena will become very marked when several strains are inoculated together, but even then the odor is not as foul as in the case of

Bacteriological Studies on Freshness of Whale Meat

69

actually putrefied fish. Also, in case of actual putrefaction proteus bacteria
are sometimes detected, but in our cases there was no such ad.vanced caseof putrefaction, and the Micrococcus flavus, M. pikowskyi, M. halophilus,
Strept. faecalis, Flavobacterium sewanense and Achromobacter halophilum
found in our examinations coincided with those detected by other researchers.
Table 4.

Species of Bacteria· detected

Name of Bacteria

Number ofstrains

Blood

Micrococcus epimetheus
Achromobacter ubiquitum2
6

rubber tree
·soil, water

3
2
1
1

MuscleMicrococcus sphero,ides
Micrococcus piltonensis
Micrococcus flavus
Micrococcus subcitreus
Micrococcus saccatus
Micrococcus halophilus
Streptococcus faecalis
Flavobacterium sewanense
Achromobacter ubiquitum
Achromobacter halophilum
Achromobacter lacticum

8
1
1

manure, soil
manure, soil
air, milk
air, water
mucous of nosesea water
animal faeces
sea watersoil, watersea water
cow's milk

1
1
1
1
1
1
7
1

.manure, soil
rubber tree
sea water
cheesehalibut
sea water•soil, watersea water

2
6

sea water
soil

Samples
examined

Intestines

Sea watet

Micrococcus piltonensis
Micrococcus epimetheus
Micrococcus pikowskyi
Micrococcus luteolus
Flavobacterium fucatum
Flavobacterium sewanense
Achromobacter ubiquitum
Achromobacter halophilum

1
3
5
.1

I
I
I

I
I

I
I

Micrococcus halophilum)
Achromobacter cystinovorum

Place ordinarily found,according to references

Conclusions. 1. Freshness is decreased in proportion to the time elapsed from catching
to dissection and to the number of bacteria detected in blood and muscle.
2. As shown in Table 4, 15 species of bacteria were detected in whale's
blood, muscle and intestines and 2 species were found in sea water. Of
the above, 6 species coincides with those reported by other re~earchers.
3. There are 4 possible routes for entry of bacteria into blood and'·muscle and it may be assumed that the intestinal tract is the most impor-70

Tomoichiro Akiba, Takeshi Tsuchiya, Makoto Umehara and Yoshiharu Katsume

tant route.
4. Although the bacteria detected may not play the .leading role inputrefaction, it is obvious that they contribute to the decrease in freshness.
5. Therefore, the most effective method to prevent decrease in freshness
is to dissect and dispose of it as soon as possible after catching.
6. ·That the previous method of salting and freezing after dissection iseffective for prevention of putrefaction, can be proven by our experimental
results on arresting increase of isolated bacteria.
References
Hunter, A. C.: Bacterial groups in decomposing salmon, J. of Bact. Vol. 5, 1920.
Jensen, L. B.: Microbiological problems in the preservation of meats. Bact. Review, Vol. 8, Nr.
3, Sept. 1944.
Sanborn, J. R. : Certain relationships of marine bacteria to the decomposition of fish. J. of Bact.
Vol. 19, 1930.
Takarada Michinori : Research on intestinal bacteria of fresh water fish, J. of Chiba Medical
Society, Vol. 20, No. 4.
Takayama Naokiyo: Bacteriological research on decomposition of lobster, J. of Chiba Medical
Society, Vol. 21, No. 9, 1943.
Weinzirl, J. : Concerning the relation of the bacterial count to the putrefaction of meat, Amer,

J. of Public Health, Vol. 14, 1924.
ZoBell, c. E.: Studies on marine bacteria. J. of Marine Research. Vol. 4. Nr. 1. 1941.

Protein Digestive Power of Sperm Whale
Pancreatic Enzyme. II.
"'Yasutaro Ishikawa
I

In the lst report, protein digestive power of pancreatic enzyme from
sperm whales was compared with that of a cow and was found much
weaker than the latter. Casein was used as the substrate and the digestive powers were measured by the method of Michaelis given in the, Japanese Pharmacopoeia.
Even after being activated in various manners, enzyme ·from whales
digested only 2.5 times of casein of its own weight while that from a cow
digested 125 times. This large difference was almost unbelievable.
However in the experiment previously reported the samples werecollected from too few bodies to discuss generally. And the time-lapse
from the caching to the dismembering of these whales was so long underhigher temperature of the summer, that there was a suspision that a partof proteases may have been destroyed by autolysis.
The present experiment was undertaken in order to eliminate these
suspicions and .to determine the true protein digestive power of a sperm
whale- pancreatic enzyme. The samples, therefore, were collected at thecoldest season during whaling (December_ 1948) and only the very freshones were used. It had been desired to collect s.amples from as many
animals as possible in order to eliminate individual differences and samples were collected from six fresh sperm whales during the two weeks at
the landstation. Ayukawa, Miyagi Prf.
EXPERIMENTAL
1) Whales Used
,

No.

Kind

Sex

Body length(ft.)

Time elapsed from·catching to dismember (hrs.)

Freshness

1

Sperm
Sperm
Sperm
Sperm
Sperm
Sperm

male
male
male
male
male
male

45
46
46
50
42
51

25
25
23
18
22
20

85
85
90
85
85
85

2
3
4
5
6

%

The freshness is the value evaluated by the whaling experts on thestation by macro-observations with considerations to the elasticity and color of

Yasutaro Ishikawa

72

the whale meat and others. The freshest state, i. e. that right after death,is taken as 100 and deterioration from that is shown by a rough percentage. Various scientific methods for the determination of freshness havebeen forwarded but none is compl~ted and procedures are too complicatedso that it has to rely on these 'empirical method.
2) Treatment a,nd Preparation of Samples
As soon as the whale body has been cut open, the pancreas taken out,
adipose and connective tissues removed as much as possible· and mincedby a hand chopper, From this were preparad 3 kinds of samples forreasons as described in (3). These samples were:a) One dried immediately with acetone.
b) ·The minced pancreas left standing overnight at room temperature
with small amount of toluen in a flask, stoppered, which was thendried with acetone.
c) The minced pancreas mixed with 30% aqueous acetone solution in
an equal amount and stored.
Samples (a) and (b) were prepared by adding anhydrous acetone tominced pancreas in an amount about 5 times its volume, well stirred andthe supernatant solution decanted. The same amount of acetone is thenadded and filtered With suction. The residue on the funnel' is washed, firstwith acetone and then with ether, and is dried by spreading it out in theroom. This is then well dried in a dessicator over Sulfuric ac:d. Procedures
up to this point were carried out at the statiori and this was ,then brought
to the laboratory for testing. The dried pancreas was first powdered bya boat-shaped mortar; sieved through a 60 mesh sieve and divided into
powders of gland substances and connective tissues. Special note should
Table I.
No.

·powder (g)Sieve residue (g)Amount sieved (%)

IA
2A
3A
4A
4B
5A
5B
6A

2.8
1.9
3.0
3.2

5.4
11.6
10.7
9.4
4.0
9.0
5.1
7.5

34
14
22
25
26
37
55
41

1.4

5.3
6.3
5.3

A denotes samples dried immediately
B denotes samples dried after being left overnight.

. .-•.!.~

Protein Digestive Power of Sperm Whale Pancreatic Enzyme. ll.

abe taken of the f(tct that the pancreas ofsperm whale contains muchlarger percentage of connective tissues compared to that of bovines, hogs
and baleen whales, so that the yield of gland substance is very poor. The
resultant yield of sieving is given in Table I.
The samples immersed in 30% acetone were brought back to the laboratory and stored at room temperature. And the supernatant solution
which might contain protease, was taken from time to time to test thedigestive power.
The kinds of samples prepared were as follow :No. 1-A,
No. 4-A,B
c
No. 2-'A,
B,
No. 5-A,cc
No. 3-A,
No. 6-A, ·B (lost), Cc
A; denotes samples dried immediately,B, samples dried after being left overnight, andC, samples immersed in 30% acetone.

Method of Activation
As in other kind of animals, pancreatic protease from sperm whales
do not show digestive action in its fresh state. It follows, therfore, that
this protease must in some way be activated and the subsequent testingof digestive powers must be carried out at the same degree of activation,in order that a compaison can be made ; if possible, at -a maximal activatedstate. For this reason, activation was carried out in following three ways
and the digestive powers determined and compared.
a) The pancreas was dried in the most fresh condition, powdered as has
been described before and this was activated by enterokinase from asperm whale.
b) Minced pancreas was left overnight at a room temperature and treated in the same way as before. This was then activated with enterokinase.
The above two were made in order to find the effect of autolysis in
the fresh condition.
c) Minced pancreas was dispersed in the same amount of 30% aqueous
acetone solution. and its supernatant solution, assumed to contain protease from the pancreas, was used as a sample for testing.
The reason that this experiment was carried out was that Kleiner and
Tauber reports2) that the protease from hog pancreas is best extractedwith 33% alcohol and is well activated by a long-time storage.
Tejima,3)3)

74

Yasutaro Ishikawa

also obtained a successful result by extracting protease from a sei whale
pancreas by the use of 30% acetone.
4) Meth0,d of Digestive Power Determination
The method of determining digestive power followed the same one as
described in the previous report1>, i. e. the Item 6 of the Japanese Pharmacopoeia for pancreatin which uses Michaelis' method of protease testing.
The substrate used was casein purified by Hammesten's method andstored in a dessicator over sulfuric acid. 5 cc. each of 0.2% solution ofthis casein (containing 2 cc. 1/10 N-KOH in 100 cc. of this solution) is taken
in test tubes and these tubes are stood in a row. A definite amount of
enzyme is dispersed in a definite ~quantity of water to extract the enzyme
and a definite amount of this enzymatic solution is added step-wise to thecasein solution and the total volume is then brought to 10 cc. each. Thissolution is kept for 1 hour at 40° to effect digestion of casein. After exactly
1 hour, 3 drops of a mixture of 1 cc. glacial acetic acid, 9 cc. water and 10
cc. alcohol is added to such test tube. One that remains clear is taken as
digested and the one showing least opalescence is taken as the limit ofdigestion. Digestive power of enzym-containing powder is found from theamount of the powder used to digest 5 cc. of 0.2% casein solution (i. e.
0.01 gm. casein) in the limit case and is then shown by the ratio of caseinto the enzymatic powder.
In the case of 30% acetone extracted solution, a definit of its supernatant solution is taken, suitably diluted and used in determination as
above.
5) Results of the Experimentsi} Acetone dried powdera) Digestive power when enterokinase is not added
Digestive power was determined with No. lA, No. 2A and No. 4B.
None of these had the power to digest casein equal to their own weight.
It follows, therefore, that the acetone dried powder either lacks digestive
power totally or has a very weak action.- This in turn may mean that the
enzyme is in an inactivated state due to freshness of pancreas or thatenzyme had been destroyed altogether. 'b) Digestive power when enterokinase is added
0.2 g. each dried powder of No. lA and No. 2A was taken, 0.05 g. ente·rokinase added end dispersed in 100 cc. of water. This was warmed at 40°

Protein Digestive Power of Sperm Whale Pancreatic Enzyme. II.

75

for 15 minutes, 30 minutes and 1 hour and activated. The result of thedetermination of digestive power using these extracts was as follows :No. lA (30min.)Digest 10 times its weight of casein but
cannot digest 12.5 times its weight.
ditto

No. 1 A (15 min.)No. 1 A (1 hour)ditto

No. 2 A (15 min.)ditto

No. 2 A (30 min.)ditto
Digests 7 times its weight of casein but
cannot digest 10 times its weight.

No. 2 A (1 hour)

From these results, it can be seen that the time of warming for activation does not differ much between 15 minutes and 1 hour but is slightly
inferior at 1 hour.
Therefore, the conditions . were set for an addition of a 1/4 amount ofenterokinase and warming for 30 minutes at 40° to effect proper activation~
The result of determination of digestive power of various samples were as
follows:
No.1 A

Digests 10 times its own weight but not 12.5 times.

No. 2 Aditto·

No. 3 A

ditto

No. 4 A

ditto

No 4 B

Digests 8.3 times iots own weigh~ but not 10 times.

No. 5 A

ditto

No. 5 B

Digests 10 times its own weight but not 12.5 times.
Digests 8.3 times its own weight but not 10 times.

No. 6 A

Digestive power of each sample made from above results is shown in
Table II, which indicates that there are hardly any difference between thesamples:
Table II. Digestive Power of Acetone Dreiecl. Powder 'Sample No.

Digestive Power(cas./powder)

Sample No.

Digestive Power(cas./powder)

IA

10

2A
3A
4A

10

4B
5A
5B
6A

8
8
10
8

10
10

'

30% Acetone Extracted Solution
The solution was stored at room temperature for about 2.5 months
during which a definite amount of the clear, supernatant solution was taken
and ·its digestive power determined. Table III, shows the relationshipbetween the digestive power and the time lapsed after immersion in acetone.
The figures denote the amount of acetone solution (in cc.) necessary toii)•Yasutaro Ishikawa

76

digest 0.01 g. of casein (shown by the least opalescence manifested) so thatthe smaller the figures, greater the digestive power.
Table Ill. Relationship between digestive power and lapse of time.
Lapse of time(days)

1 c

5
7
9

1/100

11

1/100

13
15
17
21
23
26
34
54
75

2 c

3 c

SC

6C

2/100

6/100

1/100
1/100

10/100

1/100

4 1100

1/100

2/100

1/100
11100
0.8/100
11100

1/100
1/100
0.8/100
1.5/100

15/100
1/100

3/100

15/100

1/100
1/100
1/100
1/100
1.5/100

2/100
1/100
1/100
1/100
1.5/100

12/10012/100
9/100
9/100
9/100

As can · be seen from Table III, each sample differs greatly in its
digestive power at t?e start of experiment but after about one month, allsamples, except No. 3 C, show constant vaiues of 1/100. This level is kept
until about 2 _months have elapsed after whi~h the power gradually decreases·slightly.
DISCUSSIONS
1) Direct comparison cannot be made between the value of 8-10 fordigestive power obtained by the determination of dried powder and thevalues obtained with 30% aceton solution.
However, if the amount ofwater included in minaed pancreas is considered and if the whole amountof enzyme present in the pancreas were to have transferred to the solution,then in the case of 30% acetone solution, the digestive power of dry pancreas as a whole can easily be calculated.
If the amount of the dried matter is assumed to be 25% of the mincedpancreas, and since the minced pancreas was added with an equal amountof 30% acetone, then 1 g. minced pancreas + 1 cc. ~0% acetone· . 0.25 gdry pancreas + 1.75 g extracted solution. i. e. 0.25 g dried matter froni
pancreas should have the same digestive power as 1.75 g. of the extracted
solution. Therefore, 1/100 cc of this extracted solution is equal to1
0·25. d ma tt er of pancreas.
1.
x
g dne
75 100Protein Digestive Powar of Sperm Whale Pancreatic Enzyme. II.

77

Since this amount had digested 0.01 g. of casein, then the digestive powerof the dried matter of pancreas is/ 0.251
70·01 '1.75
xToo =

i. e., it has digested 7 times its own weight of casein. Of course, this .
calculated amount shows the digestive power of pancreas as a whole but inthe case of acetone dried power, it has been put through the sieve during
which purification of the enzyme may have been efected so that a precise
comparison cannot be made as to which is larger. However, it is consideredthat the order is well adapted in this case.
2) When enterokinase is not added to the dried powder of pancreas,· nodigestion occurred. From this fact, it can be assumed that these pancreas
was stored in a comparatively fresh state until treatment and that protease
had not been destroyed and stayed in its natural state.
The digestive power of the dried powder of pancreas when activatedby the addition of enterokinase was found to be about the same in eachindividual and digested 8_:__10 times its weight of casein. The protein
digestive power of 30% acetone extracted solution varies greatly at first
but becomes most activated about one month after immersion at whichtime, with the exception of one sample, the values are exactly the samefor all the samples. Ih this oi1e exception, i. e. 3 C the supernatant solution
was found to be considerably opalescent due to the inclus:on of grease (theothers were almost clear). This is apparently the error due to the inclusionof a large amount of adipose tissues at the time of sampling.. The value
when this same level had been reached in all the sample, i. e. 1/100, showsthat the dry matter of the original pancreas would digest ~bout 7 times
its own weight of casein.
From these results, it is considered that the digestive power of proteasefrom sperm whale is considerably inferior to that from bovines and hogs.
However, this is true only when casein has been used as a substrate andit is doubtful whether the same tendency will be manifested when otherproteins are chosen as a substrate.
From the results of. these experiments, it can be assumed that there
is very little difference between individuals.

78

Yasutaro Ishikawa

SUMMARY
Protein digestive power of pancreas from 6 sperm whales 'Yas determined and following results were obtained :1) Acetone dried powder does not show digestive action as it is.
2) When acetone dried powder is activated by the addition of 1/4 its
amount of enterokinase from a sperm whale and warmed for 30 minutes
at 40°C, it digests 8-10 times its own weight of casein and there were
no differenc between individual animals.
3) 30% Acetone extracted solution of minced pancreas, reached its maximal activated state after storage of about one month at room temperatureand the digestive power was the same for all the samples, i. e. 1/100 cc.
of this. solution digested 0.01 g. of casein. Calculated from this value, thedry matter of the pancreas used would have digested about 7 times its
weight of casein.
References·1) Ishikawa: Bulletin of the Japanese Society of scientific Fisheries Vol. 14, No, 2, 107 (1948);
Ishikawa and Tejima: Sc. Report of the Whale Reserch Institute, No. 2, (1949).
2) I. S. Kleiner and H. Tauber: J. Biol. Chem., 104, 267 (1934).
3) Tejima a1t1d Ishikawa: Lecture read before the 3 rd Meeting of the Whale Reserch Institute in October, 1948, unpublished.

Properties of Fats and Oils contained in Various
Parts of a Sperm vVhale Body .
Takajiro Mori and Masamichi Saiki
Introduction
.

1-33)

Numerous studies have been made since the olden times on whale oil.
Many works have been mad~ . on its physico-chemical properties, on thecomponents of its fatty acids and its saponifiable matter and their properties. However, these studies have chiefly been confined to the oils fromwhale blubber, bones and head cavity and there has been very little evidence of chemical studies made on oils from large and small intestines,
liver, kidney and other organs. It seemed that it would not only be interesting to make a study of oils from various organs from , the point of fat
metabolism but also necessary from the point of their utilization. Therefore, studies in this line were made.
Experimental materials
The present experiments were carried out with 2 heads of sperm
whales (Physeter macrocePhalus L.) which were caught off Kinkazan point(Miyagi Prefecture in the northeastern part of Japan). One was a male(Sperm Whale B). measuring 34 feet long, caught in September 1946, andthe other, also a male (Sperm Whale A), measuring 35 feet long, caug~t
on 23rd. September 1947. Each individual animal was dissected in thedissecting room, various parts collected and those from which oil can beobtained by boiling, i. e; skins, large and small intestines, were boiled on the
spot. Those difficult to put to this procedure, i. e. liver and large and small
intestines of the Sperm Whale A, were frozen and sent to the laboratory.
In the laboratory, these organs were defrosted, dehydrated by treatment
with alcohol, car.bon dioxide then blown in and dried in· vacuum. After
this dried organs were pulverized, it wa~ extracted by ether in a Soxhlet
The alcohol used for dehydration was evaporated under
extractor.
reduced pressure and dried and this residue was shaken with ether. This
ether and that from the Soxhlet extractor were brought together and after
drying with anhydrous sodium sulfate, the ether solution was dropped into

Takajiro Mori and Masamichi Saiki

80

dry· acetone while ·stirring constantly and the phospholipid precipitated
was filtered off.34>
Acetone and ether were driven off under reduced pressure in carbon
dioxide stream and some oil was obtained. The ether extract from theliver of Sperm Whale B amounted to 11.73% of dried matter. The oilfrom the head cavity was collected when the oil flowed out at the time ofdissection. This head oil (oil from the head cavity) is a white solid at aroom temperature, forms a pale, yellow liquid at (around) 30° and precipitates about half the amount of white, clou4y crystals at around 25°.
The blubber oil ( Oil from the skin) is pale yellow. The oil obtained
from large and small intestines is yellow, but that obtained by ether
extraction is brown. During the summer, intestinal oil is liquid but pricipitates solid matter during the winter (11°C). Liver oil is brown andremains liquid, although slightlY viscous, all through the winter.
These samples were treated w:[th acid clay or activated charcoal andyellow oils were obtained which were used for the following experiments.
, Experiment
Neutral oils- With the above samples, specific gravity, coefficient ofrefraction, add value and saponification value were measured in an ordinarymanner, and the ipdine value by Wijs' method. Saponification was limited
to 2 hours. Vitamin A content was also measured by Beckmann's spectrophotometer. The results obtained are shown in Tables I and II.
Neutral Oils from Sperm Whale A.

Table I.

ri

Non40° Acid
Vitamin
Sapon. Iodine
nD
Value Value Value sapon. A (I. U.)
matter- - -- - - ---1--Head oil
White soild
140.71 ! 55.01
0.8744 1.4500
0.73
44.98
103
·BlubberPale yellow liquid
0.8779 1.4570
0.75
131.08
82.33
35.!?1
370
Small intestine Pale yellow liquid
0.8879 1.4570
2.01
156.84
78.88
17.35
865
Large intestine Pale yellow liquid
1.4522154.48
13.36
77.71
19.18
1
Liver
[Brown, viscous liquid
1.4711
2.64
165.3492.41
19.01 12,655
Par1;sAppearance

Parts

Appearance

25°

d o
4

Table. II.

.Head oilBlubber
Small intestine
Large inte.stine
Liver

I

White solid
Pale yellow liquid
Pale yellow liquid
Pale yellow liquid
Dark brown
semi-solidI

Neutral Oils from Sperm Whale B.
26"
32°
Acid Value I Sapon.
I Iodined 4onD! ValueValuer0.87520.87920.90560.9054-

1.46891.46961.4721
1.47241.4861

0.52
0.36
1.03
1.01
18.83

142.35127.89155.91

I

57.57
82.13
78.23
79.41
82.12

Properties of Fats and Oils contained in Various Parts of a Sperm Whale Body

. 81

Although the whales A and B are different individuals," all values inthe two above tables are similar, showing that there are very little difference between the individual animal.
Non-saponifiable matter- The samples were saponified with alcoholic potash,
water added and shaken 4 times with ether. combined ether solution waswashed with water, dried with sodium sulfate and ether evaporated underblowing in carbon dioxid~.
Non-saponifiable matter from all the samples were a pale yellow solid
during the winter (11°C) and the properties are as shown in Table III.
·The melting points were determined in a capillary tube, values being thatof the beginning and end of melting.
Tabfo ill.
Parts

Properties of Non-Saponifiable Matter from the Sperm Whale A.
Appearance

Head oilBlubber
Small intestine
Large intestine

I

Pale yellow solid
Pale yellow solid
I
Pale yellow solid I
Pale yellow solid

40Q
nD

I

1.44501.45001.4490I

1.4500II
I
I

Melting point

Iodine value(Wijs')

35.0-37.0
20.5-24.0
29.5-31.8
29.D-31.8

39.08
71.74
51.72

(QC)

50.53

Fatty acids- The aqeous solution obtained by the removal of nonsaponifiable
matter was evaporate'd under reduced pressure to remove alcohol, hydrochloric acid added and ·the freed fatty acid extracted by ether. This was treatedin the same way as for non-saponifiable matter. Of the fatty acids obtained:
Mixed fatty acids from the skin oil was a pale brown liquid during thewinter (11°C) precipitating some solid fats ; That from head oil was paleyellow solid, with a mixture of some crystalline solid fats ; and those fromlarge and small intestines were a yellow solid. Properties of these fatty
acids are shown in Table IV.
Table IV.

Mixd Fatty Acids from the Sperm Whale A.

Parts

40Q
nD

Melting point(QC)

lodine value(Wijs')

Head oilBlubber oil
Small intestine oil
Large intestine oil

1.44371.45111.45301.451816.0-20.0
13.5-18.0
28.0-31.0
28.0-31.5

56.44

246.68

80.57
79.54

199.22194.66

81.85

195.:n'I

I

I

Neutralization·
Value

Solid and Liquid Acids- Solid and liquid fats were obtained from the
mixture of fatty acids using about 10 g of each sample according to Twitchelrs method:i.">. The solid acids here obtained were white solids, andliquid acids from intestinal. and head oils were yellow,· that from blubber

82J

Takajiro Mori and Masamichi Saiki

oil, yellowish brown. . The properties of these solid and liquid acids areshown in Tables V and VI.
Table V.
--------

Parts
Head oilBlubber·------------I!

!

Small intestine I'
Large intestine

Appearance

Solid Fatty Acids from the Sperm Whale A.

·---llagainst
ercentage \total
5fatty acidI 21.31% 'White solidWhite solidWhite solidWhite solid
Tale VI.

10.07%
18.64%
19.29%

1

Appearance

1.42891.4352
1.4363
1.4368Percentage

I

point

'

("C)

'

38-39
50-52
50-52.5
50-52.540°! against totalnDfatty acid-------1Yellow liquid
78.69%i

Head oilBlubber
Small intestine
Large intestine- ~fMelting - Iodine value Neutraliza(Wijs')

tion value

3.06
14.56
15.39.
11.48

250.63213.62211.36210.49Liquid fatty acids from the Sperm Whale A.

,.--------

Parts

60°

nD

Yellowish brownIliquidYellow liquid
I!
Yellow liquidI89.93%
81~36%

80.71%

Iodine value(Wijs')

Neutralization
value-----------

I

1.44501.45221.45501.4542

73.93
92.26
95.07
91.03

236.12194.72
187.25187.83

Summary and Discussions
1) The blubber oil of the sperm whale, especially the oil from its headcavity, is composed of fatty acidsof small molecular weight and is unique'
to it except the butter fat. As a result of the present experiments, it has
been found that the average molecular weight of fatty acids was thesmallest in those contained in head oil, that from the blubber oil next andfollowed by large and small intestine and liver oil, which contained thelarge molecular weight acids although the values were approximately similar(estimated_ from the amount of non-asponifiable matter of neutral fats andthe saponification value).
Both the solid and l:quid acids from the head oil were of small molecular weight compared to other oils from other organs.
2) The degree of unsaturation of mixed fatty acid is the smallest in thehead oil, those from blubber, large and small intestines are about the sameand are larger. This seems to be due to the fact that the head oil contains a large amount of solid acids and that the degree of unsaturation ofits liquid acid is small compared to the oils of other organs.
3) It is known that sperm whales feast largely on quantities of squids
proof of which was borne out by the discovery of a large amount of themin the stomachs of the sperm whales used for these experiments. In order.

..

Properties of Fats and Oils contained in Various Parts of a Sperm Whale Body83

to find the relationship between the specific nature of sperm whales andthe oil of squids devoured as their chief food, reference was made to theanalysis of the oil from a certain kind of squids caught in the Hokkaido,
Japan, by Mitsumaru Tsujimoto:IB\This analysis showed that this oil
contained a very small amount of non-saponifiable matter and its degree ofurrsaturation was exceedingly high. Comparison of similar data by E.
Andre and H. Canal37) showed that the degree of unsaturation was alsohigh in the French squid oil but contained more non-saponifiable matterthan the Japanese squid oil, values being near those of sperm oil. Accor·
ding to these French investigators, the non-saponifiable matter contains,differing from the sperm oil, no aliphatic alcohols, either saturated or
unsaturated. From these facts, it is assumed that sperm whales utilize
these squid oils to turn into its own oil.
4) As has been shown in the foregoing, sperm whales possess a large
amount of wax so that it gives abnormally high amount of non-saponifiable
matter. In the present experiments, the amount of non-saponifiable matterin the head and bfobber oil was exceedingly large but that from large andsmall intestine and liver oil was only one-half of the former, being about
the medium between the oil of sulfur bottom and that of sperm whale. It
is interesting to note that the average moleclar weight of fatty acids andthe amount of non-saponifiable matter are in an inverse ratio in the abovetwo.
It would be interesting to study the relationship between these facts
and the fat metabolism in whale bodies and if such a relation does exist,
what significance it has. These must be left for the future.
5) The degree of unsaturation of the non-saponifiable matter is the smallest in head oil, followed by the oils of large and small intestines whichwere about the same, and largest in the blubber oil.
Our thanks are due to the Department of Education for grants in
aid of this research.
References
1) Mitsumaru Tsujimoto: J. Ind. Chem., Japan. 13, 1267-1246 (1910).
2) Mitsumaru Tsujimoto: J. Tokyo Ind. Lab., 15 (10), 1-80 (1920).
3) Mitsumaru Tsujimoto: J. Ind, Chem., Japan. 20, 41-44 (1921).

84

Takajiro Mori and Masamichi Saiki

4) Mitsumaru Tsujimoto: Chem. Um3chau. 30, 33-36 (1923); J. Ind. Chem, Japan. 26, 60862J ; Tokyo Ind. Lab., 18 (2), 103-123 (1923 ).
5) Mitsumaru Tsujimoto:Chem. ·um-,chau. 32, 127-8 (1925).

6) Mitsumaru Tsujimoto:Chem. Um>chau. Fette. Oele. Wache und Harze, 32, 202-4 (1925).

7) Mitsumaru Tsujimoto and Hosuke Kimura: Tokyo Ind. Lab., 22 (12;, 53--63 (1927).
8) Mitsumaru Tsujimoto: Tokyo Ind. Lab., 23 (3), 61-63 (1928).
9) Mitsumaru Tsujimoto and Hanji Koyanagi: J. Ind. Chem., Japan. 40, 391-395 (1937).
10) Mit3umaru Tsujimoto and Hanji Koyanagi:

J. Ind. Chem., Japan. 40. 694-696 (1937).

11) Emile Andre and Th. Francois: Compt. rend. 183, £63-6c5 (1926).
12) Emile Andre and Th. Francois:

Compt. rend. 185, 279-281 (1928).

13) Emile Andre and Th. Francois: Bull. Soc. Chim. biol. 9, 117-125 (14) Emile Andre And Th. Francois:15) Shushi Toyama:).

Compt. rend. 182, 497-499 (1925).

Tokyo Ind. Lab., 22 (11 \ 225-227 (1927).

16) Shushi Toyama: J. Ind. Chem.,.Japan. 30, 519-·527 (
224 (1927).

) ; Tokyo Ind. Lab., 22 (11), 189-

17) Shushi Toyama: J. Ind. Chem. 30, 527-532 (1927).
18) Shushi Toyama and Chitaro Tsuchiya: J. Japan Chem. Soc. 56, 1072-1077 (1935).
19) Shushi Toyama and Chitaro Tsuchiya : lst Report. J. Japan Chem. Soc., 56, 105')-1054 (1935);
2nd Report. ibid., 56, 1055-1059 (1935).
20) Shushi Toyama and Chitaro Tsuchiya: J. Ind. Chem., Japan. 38, 627-631 (1935).
21) Shushi Toyama and Goroku Toyama: J. Japan Chem. Soc., 56, 1316-1319 (1935).
22) Shushi Toyama and Goroku Toyama: J. Japan Chem. Soc., 56, 1077-1080 (1935).
23) Shushi Toyama and Chitaro Tsuchiya: J. Japan Chem. Soc., 56, 1313-1315 (1935).
24) Shushi Toyama: J. Japan Chern. Soc., 59, 810-814 (1938).
25) Seiichi Uyeno: J. Ind. Chem., Japan. 38, 1349-1344 (1935).
26) Seiichi Uyeno and Ryosei Koyama: Bull. Chem. Soc. Japan, 11, 394-4fl3 (1936).
27) Seiichi Uyeno and Ryosei Koyama:28) Seiichi Uyeno:

J. Japan Chem. Soc., 57, 1-9 (1936).

J. Ind. Chem. Japan., 41, 643 (1938).

29) Koichiro Ura: J. Ind. Chem. Japan, 11, 854-864 (19G8).
30) Shigetada Tanimura: · J. Marine Prod. Inst., 15 (2), 37-54 (1920).
31) T. P. Hilditch and J. A. Lovern: J. Soc. Chem. Ind. 47, 105-111 (1928); ibid., 48, 359-365;
ibid., 48, 365-368 (1929).
32) S. Schmidt·Nielson and A. Flood:

Kg!. Norske Viden~ Kab. Scks Kab. Foresh. 5, 198-201(1932).
33) Tei Ando, Sukenori Uzawa and Isao Shigetomi: J. Japan Mar. Prod. Assoc., 10, 1, 17 (1941).
34) W. R. Bloor and Ruth H. Snider: J. Biol. Chem. 87, 399.
35) E. Twitchell: J." Ind. Eng. Chem, 13, 9, 806-807 (.

).

36) Mitsumaru Tsujimoto and Hosuke Kimura: J. Ind. Chem., Japan. 30, 865 (1927); Cf. Tsujimoto:37)

J. Ind. Chem., Japaan. 17, 2Jl, 1300 (1914).

E. Andre and H. Canal: Compt. rend. 183, 152 (1926).

Studies on kitol 1.
Preparation of Kitol from \Vhale liver Oil
Tadashi Tawara and Ryusuke Fukazawa
Willstaedt & Jensenl) and Nakamiya2l pointed out the presence of a
Vitamin. A-like substance, differing from the ordinary Vitamin A in its
properties, in whale liver oil. Recently, Baxter, et al.3l isolated kitol.
The presence of a foreign substance in whale liver oil had long been
predicted by the difference of. their spectrum analysis and the Carr-Pricereaction from ordinary fish liver oil. The maximum of the absorption
spectrum of the unsaponifiable matter of ordinary fish liver oil is in theneighborhood of 328 mp, but that of whale liver oil is at 290-325 mp, andthe Carr-Price reaction of whale liver oil is blu1sh-purple whereas that offish liver oil colors· blue. From the proximity of the figures of extinctionof 265 mµ and 328 in,u in the absorption spectrum of the.
coefficient,unsaponifiable matter of whale liver oil, it had been believed that theabsorption curves would lie close together and overlap each other and an
apparent maximum would lie in intermediate point.2)i!·

The authors also determined Fig. 1 Unsaponifiable matter of fin whale liver oil
E rtthe extinction coefficient E 11.%cmof the unsaponifiable matter offin. whale liver oil (Fig. 1)* by
the spectrophotometer and wereable to prove this point.
This absorption curve apparently differs from that .of
tunny liver oil (Fig~ 2). The
greatest difference exists in theposition of the absorption maximum and the width of the curve. The Carr-Price reaction of
220 2~0 240 250 260 ~70 280
this unsaponifiable matter of .,.

* The reason that the curve is so wide is because the· absorption is so indistinct that the valueof extinction coefficient, E1 % , does not come out clearly.
1cm

Tadashi Tawara and Ryusuke Fukazawa
Fig. 2 Unsaponifiable matter of tunny liver oil

fin whale liver oil gives bluishpurple color while that of tunny
liver oil, blue.
The oxidation product of
H-+---l---+--+---1-+---'---1---i--+--'-l-+--i600
Vitamin A gives red or reddish
H-+--+--+--+--1--1---'---+--+--+---'-lr--+-15° 0
brown coloring by this reaction.
According to the authors' experiments, majority of this oxidation product transists into saponifiable matter during saponification so that it does notseem to constitute a reason for.__...__.....__,_---'-__,_.1---'---'--'---'-,..-'"''"=-=~100
bluish purple coloration. Naka220 2,0 240 250 260 210 280 290 ,co ,10· 520 ,,a >40 550mµmiya and others harbored adoubt that it might be due to the large amount of cholesterol present but
the authors take the view that this is due to the presence of kitol.
Kitol as isolated by Baxter, et al.,· were colorless, elongated prismatic
crystals of Fp. 88-90°, with a molecular formula corresponding to CoH5s(OH)2, possessing 8 double bonds. It gives an absorption maximum at
290 mµ, and E 1% 290 mµ=707.
1cm
The fact that kitol possessed twice the molecular weight of Vitamin A
was utilized in separating the two by the difference of solubility. The
unsaponifiable matter of fin Fig; 3 Unsaponifiable matter of fin whale liver oil-Fraction A

whale liver oil was dissolved in. petroleum ether and this was
extracted with 90% methanol
into which Vitamin A was transited. The substance which remained in the petroleum ether
solution to the last was examined by the spectrophotometerand the. value of E 1.%. at each.
1cm
wave length was shown by acurve (Fig. 3). This substanceshowed an absorption maximum .
2}0 240 250 260 270 280 290
at 290 mµ, and the Carr-Price 220mµ~~__,.-.~----~-~~~__,,._~~c..__,__,__,100)}0)40 )50

Studies on Kitol. I. Preparation of Kitol from Whale liver Oilreaction gives a red coloration.
Apparently, it contained a verysmall amount of Vitamin A.
On the other hand the E 1%
1cmvalue of the substance that tra-87

Fig. 4 Unsaponifiable matter of liver oil fin whale-Fraction B

nsite9 to 90% methanol (curvein Fig. 4) gave a maximum
absorption at around 328 mµ, **which is very similar to thecurve of tunny liver oil· (Fig. 2).
It does not give the indistinct
curve as in Fig. 1, any more.
The Carr-Price reaction is blue,
so that this substance must be
nothing but Vitamin A.
From these facts, it can, be assumed that the substance that reamainedin petroleum ether to the last is clearly kitol.
EXPERIMENTAL
250 g. fin whale liver oil was warmed with 500 cc of 20% methanolic
potash for 1 hour in a water bath. After saponification, this was extractedwith ether and the ethereal layer was washed several times with waterand dried with anh~drous Glauber's salt. After distilling off ether, 54 g. of
unsaponifiable matter was obtained. This substance gives bluish purple
coloration by Carr-Price reaction and, according to E 11cm.% curve (Fig. 1) byspctrophotometer* **, ,Possesses an absorption maximum 303 mµ although
the absorption area is so indistinct that it is hard to obtain a correct valueof: E11.l.%.
Accordingly, the curve became one of a wide bartd. This is appa~cm

rently due to the fact that two or more substances are mixed in it andgives a complex absorption so that this must be taken as the apparent
absorption maximum.
The .total amount (54 g) of this unsuponifiable matter was· dissolved in1000 cc. of petroleum ether (Bp. 30-60°), placed in a separating funnel** According to this experiment, the absor13tion maximum of Vitamin A-like substance isolated
from the unsaponifiable matter of tunny liver and whale liver oil is more nearer 325 m11 than
328m,11.
*** Spectrophotometer by Carl Zeiss was used. The solvent was absolute alcohol.

'

88

Tadashi Tawara and Ryusuke Fukazawa

and extracted 38 times with 500 cc each 90% methanol. The petroleum
ether layer was then washed several times with water, dried with anydrous
Glauber's salt and distilled under reduced pressure. A reddish yellow,
syrupy matter was obtained to the amount of 14 g. 90% Metanol solution
was distilled under reduced pressure in carbon dioxide stream and the'residue, composed mostly of water and a small amount of oil drops floating
on it, was placed in a separating funnel. This was extracted with ether
and the ethereal layer was· distilled off after dry in with anhydrous Na2S04.
The residual matter, reddish yellow syrup obtained was ea. 39 g. About
1 g unsuponifiable matter was lost during this separating process.
The substance which transited to 90% methanol (hereafter designatedas fraction B) gives a blue coloration to Carr-Price reaction, extinction
coefficient curve as shown in Fig. 4 which approximately coincides withthat (Fig. 2) of the unsuponifiable matter from tunny liver oil, i. e. itshows an absorption maximum at around 328 mµ and does not give theindistinct absorption as before separation. The Carr-Price reaction is notlonger bluish purple but blue so that this is a substance devoid of foreign
matter, i. e. Vitamin A alone.
The residual matter in petroleum ether "(hereafter designated fractionA) gives a red · coloratin to Carr-Price reaction and extinction coeffcient
curve (Fig. 3) ·is clearly different from that of fraction B (Fig. 4) andpossesf]es an absorption maximum at 290 mµ, Eir~ 290 mµ=500.**** It isapparent that fraction A is kitol.
No presence. of kitol were discovered. in the oil obtained from theeyeball of sperm and sei whales ..
The authors' deepest gratitude is due Dr. Tsutomu Maruyama, Directorof the institute, who gave them kind ·instructions, and to Mr. Tadashi
Nakai of this institute and to Dr. Seiichi Ishikaw~ of the University of
Literature and Science for their unfailing assistance and advice.
References
1) . H. Willstaedt and H. B. Jensen: Nature, 143, 474 (1939).
2) J. Nakamiya,_ K. Koizumi and I. Kawakami: Riken Iho (Report of the Institute for Physicaland Chemical Research), 20, 576 (1941).
3) F. B. Clough, H. M. Kascher, C. D. Rob_eson and J. G. Baxter: Science, 105, 436 (1947); M.
H. Stern, C. D. Robeson, L. Weisler and J. G. Baxter: J. Am. Chem. Soc., 69, 869 (1947).
4) Embree: J. Am. Chem. Soc., 65, 910 (1943).

**** Extinction coefficient of pure kitol at 290 m,u is 7073) but that of kitol separated by "Embree, etal.4J at first was 580 at 290 mµ, which apparently was still quite impure.
obtained by the author theoretically contains 70% kitol.

Frantion A

Studies on Kitol. II. Influence of Kitol Fraction on the
Determination of the International Unit of Vitamin A.
Tadashi Tawara and Ryusuke Fukazawa
In the previous paper*, the authors showed that the unsaponifiable

matter of whale liver oil gave only the apparent absorption maxin:ium dueto the presence of a mixture of two or more substances while that oftunny liver oil gave an absorption maximum at around 328 mp. The
Vitamin A fraction (fraction B) of whale liver oil gives a maximum ataround 328 mp but due to the presence of kitol fraction (fraction A), thewhale liver oil gives a very complicated absorption. Accordingly, thedetermination of Vitamin A of whale liver o:l by a spectrophotometer does
not give the correct extinction coefficient at 328 mp, only that of an apparent value, so that determination of true Vitamin A content cannot bemade by this method. This fact is borne out by the fact that with whale
liver oil, results of animal test and spectrum analysis do not coincide.'
The authors examined the influence of kitol fraction on Vitamin A by
the use of fractions A and B separated by the authors. However, this is
an intermediate report and further detailed experiments must follow since
there is no direct proof that the kitol fraction obtained does not containeven a minute amount of Vitamin A, or that it does not contain otherunsaponifiable matter.
39 g. of Vitamin A fraction and 14 g. of kitol fraction were obtained
from 53.5 g. of unsaponifiable matter of the whale liver oil. First, theinternational, unit· of Vitamin A was determined with Vitamin A fraction
and then the extinction C_?efficient at 328 mp of a mixture of 14 parts kitol
fraction to 39 parts Vitamin A fraction, which the international unit ofVitamin A was calculated. As a result, it was shown that the value ofthe international unit of Vitamin A in this case came out 12.1% higher
1than the true value of Vitamin A content. In other words, the true eontent of Vitamin A is the value obtained by subtracting 12.1% from thevalue obtained by spectrophotometric determination.
It follows, naturally, that the determined value of Vitamin A comes* Cf. Report I, p.. 85 of ·this Bulletin.

90

Tadashi Tawara and Ryusuke Fukazawa

out larger than the true content of Vitamin A when the amount of kitol
content becomes large compared to the amount of Vitamin A. This willshow that a presence of kitol has a very great effect on the assay ofVitamin A by the spectrophotometer and therefore, points to the fact thatthe method of Vitamin A determination of whale liver oil must be
improved.
The authors' thanks are due to Taiyo Fisheries Co., for their kindness
in furnishing the material for these experiments.
EXPERIMENTAL
53.5 g unsaponifiable matter were obtained from 250 g. of the liver oilof a fin whale by the method as described in the previous report. From
this unsaponifiable matter, 39 g Vitamin A fraction (fraction B) and 14 g.
kitol fraction (fraction A) were obtained (loss of 0.5 g.).
1) Carl Zeiss' spectrophotometer was used to determine the international
unit of Vitamin A fraction (solvent-absolute alcohol).
1% 328m 11=454.5·B fraction-E 1cm,.
therefore, this substance contains 727,000 I. U. of Vitamin A per 1 g.
2) Kitol fraction (fraction A)-E 11cm
% 290 mp=500
E 1lcm% 328mr1i=108.7
therefore, this substance would look as though it containnd 174,000 I. U. of
Vit~min A per 1 g., but the Carr-Price reaction gives a red coloration andcontains virtually no Vitamin A.
3) International unit was determined with a mixtue of 39 parts Vitamin
A· fraction and 14 part kitol fraction (a pr0portion so arrived because theunspo11i:Wible matter of this whale liver oil contained these 2 fractions in
a proportion of 39 : 14). E11cm% 328mp=375·
therefore, this substance must contain 600,000 I. U. of Vitamin A.
However, 1 g. of this substance contains only 39~ 14 g. of Vitamin A
fraction and, taking that the kitol fraction does not contain any Vitamin A(by Carr-Price reaction), the true content of Vitamin A can be shown by
the following equation :727,000 I. U. x -~~ · .535,000 I. U.
therefore 600,000-535,000=65,000 I. U.
Actually, 65,000 I. U. Vitamin A had been shown than was actually con-

The Influence of Kitol Fraction on the Determination of the International Unit of Vitamin A 91

tained, or in other words 12.1% more Vitamin A.
4) The determination of internatio~al unit was mad~ with a mixture ofan equal portin of A and kitol fraction which gave :1.% 328 mµ=35.8
E.lcm
therefore,. this substance seems to contain 572,800 I. U. Vitamin A but
according to the aforementioned calculations, it actually contains only
363,500 I. U. In this case, the spectrophotometric value gave 30.4% higher
content of Vitamin A than was actually present.
5) International unit was determined with the unsaponifiable matter oftunny liver oil which gave :E1lcm.% 328 mµ= 38.5
therefore, 1 g. of this substance contains 61,600 I. U. of Vitamin A.
6) International unit was determined with a mixture of 39 parts unsaponifiable matter of tunny liver oil and 14 parts kitol fraction· which gave :E1.% 328 mµ= 50.0
lcm
therefore lg. of this substance seems to contain 80,000 I. U. of Vitamin A
but according to the above calculation, the real content of Vitamin A is
45,100 I. U. In this case, ea. 77.4,o/o higher value than the actual content
of Vitamin A was obtained by the spectrophotometric determination.
7) International unit was determined of an equal amount of unsaponifiable
matter of tunny liver oil and kitol fraction which gave :E 1.% 328 mµ=·59.0
lcm
therefore 1 g. of this substance shows a Vitamin A content of 94,400 I. U.
but, according to foregoing calculation, the actual content of Vitamin A
would be 30,800 I. U., if there is no presence of Vitamin A in kitol fraction.
In this case, the photometric determination has given 206.5% higher value
than the true content of Vitamin A.

·.

Studies on Kitol. II1. The Effect of Sunlight,
Air and Heat on the Vitamin A and Kitol Fractions
Tadashi Tawara and Ryusuke Fukazawa
In the 1 st Report, the separation of Vitamin A and kitol fractions

from unsaponifiable ip.atter of a whale liver oil was described. In order torefine kitol, it became necessary to know its properties, chiefly its stability,
and the effect of sunlight, air and heat on these two fractions were
examined.
1) The sunlight has a. considerable effect on both fractions. Direct radiation of sunlight on 0.1% alco]J.olic solution of Vitamin A fraction destroyed
one-half the amount of Vitamin A in 3 hours and that on kitol fraction
destroyed one-half of it in 4 hours.
2) Continuous passage of air through 0.1% alcoholic solution of kitol fraction showed no marked effect on kitol but the same process on Vitamin A
fraction resulted in the destruction of about 2/3 the amount of Vitamin A
present after 5 hours.
3) Boiling 0.1% alcoholic solution of kitol did not show any damage tokitol in 12 hours but the same treatment on Vitamin A fraction resulted
in the destruction of ea. li7 the amount of Vitamin A.
As a result, it can be stated that Vitami A is very unstable toward·sun light and air, and is slightly effected by heat. Kitol, on the other
hand, is fairly stable against heat and oxygen but is considerably sensitive
to light.
EXPERIMENTAL
The Vitamin A and kitol fraction obtained by the method as describedm Report I were dissolved in absolute alcohol to obtain a 1% solution,
respectively. With this solution, following experiments were carried out
the amount of Vitamin A and kitol remaining in the respective solutions
were determined by measuring the extinction coefficient at 290 mµ for kitol
and at 328 mµ for Vitamin A.

1) The Effect of Light
30 cc. each 0.1% alcoholic solution of Vitamin A and kitol fractions were

The Effect of Sunlight, Air and Heat on the Vitamin A and Kitol Fractions

93

placed in 5 graduated test tubes (dia. ea. 1.3 cm, length ea 21.5 cm; co!orless, hard glass) each and after being stoppered, were radiated to direct
sunlight (on 6th September 1949. Fine. From 1200 to 1700). One test tube
each of Vitamin A and kitol fractions were taken every one hour and thevalues were determined. After radiation was completed, the content of thetest tuve was diluted with absolute alcohol to bring the concentration to
0.002% and the extinction coeffecient was measured with Carl Zeiss'
spectrophotometer, that for Vitamin A fraction at 328 m11, and for kitol
fraction at 290 mp. The results of the measurements are shown in Table
I and Fig. A.
Table I.

The Influence of Sunlight(Vitamin A Fraction)

0 hr.

Radiation time

1 hr.

2 hrs.

3 hrs.

I
I
I

El% 328mµ
lcm

417

312

278

228

--------I

I

I

4 hrs.

I

.156

j
I

5 hrs.
125(Kitol Fraction)--f%--0.

E lcm290mµ

500

417

278

385
Fig

243

192

A

600

E 1lcm% 328mt'or
E 1% 290mu

l=

I ""'
400

500

''

'~

Kitol fraction'' ',...............

200

2

.. .

"'"

" ..........

'

4

100

0

Vitamin A fraction

---

500

5

---houre

2)

The Effect of Heat (78°C)
500 cc. each of 0.1% .alcoholic solutions of both fractions were taken
into l /. Erlenmeyer flask and boiled with a reflux co:1denser in a boilingwater bath (78°C). After each 2 hours period, the solution was slightly
cooled and 30 cc. of .it was drawn out for testing. This was diluted to
0.002% strength with absolute alcohol and determinations made with
spectrophotometer. Results are showri in Table II and Fig. B.

94

Tadashi Tawara and Ryusuke Fukazawa
Table IT.

The Influence of Heat (78°C)(Vitamin A Fraction)

Heating time

0 hr.

2 hrs.

4 hrs.

E1% 328mµ
!cm

417

417

385

I 6 hrs. I 12 hrs.
I

360

360

500

500

(Kitol Fraction)
E 1 % 290mµ
!cm

500

500

500

Fig. B600

E 1 % 328mµ
!cm

'°°

or
E1 % 290mµ
lcm

400

I

Vitamin A fraction---------------·~

Kitol fraction,...__

,oo
200

100
2

0

4

6

8

10

12

houre

3) The Effect of Air
15 cc. each of 0.1% alcoholic· solution of both fractions were taken in
the graduated test tubes (as described above), and the air was passed
through this solution. of the extent of its surface is continueing constantly
noise by means of a glass tube measuring ea. 0.5 mm. in inside diameter.
This procedure was carried out from l' to 5 hours. When the passage
was stopped, each glass tube was washed with little alcohol and thiswashing was added into the glass tube and the volume brought correctlywith absolute alcohol to 15 cc again (alcohol evaporated ea. 1.5 cc. per hour
during passage of air). Absolute alcohol was further added to thi~ solution
to make it a 0.002% solution and the value were deterlll,ined as previously.
Results are shown in Table ill and Fig. C.
Table ill.

The Influence of Air(Vitamin A Fraction)
Passage time

0 hr.

. E1% 328mµ
lcm

417

I
I

1 hr.
417

I

2 hrs.

3 hrs•

4 hrs.

385

360

333

455

455

5 hrs.

I

125(Kitol Fraction)
E1% 290mµ
!cm

500

500

455

455

The Effect of Sunlight, Air and Heat on tl1e Vitamin A and Kitol Fractions
Fig. C
E1lcm% 328m"ror
E1% 290mu

lcm

'

600
500
400

----

Vitamin A fraction-- - - --........

~~-·······················

Kitol fraction

~

.500

\

200

\

100
0

2

4

----hot1rs

595

On the Respiratory Pigments of Whale(Studies on vVhale Blood II.)
Tadashi Tawara
The diving method of whale is very regular and ordinarily this may
be divided into the two classes of short diving and lorig diving. That is,
it makes a relatively shallow dive, then breathes by raising its head andback. above surface of water and then makes a shallow dive again. The
diving and breathing is repeated several times. This repeated dives arecalled surface dives. After repeating this surface dives several times, it
makes a deep dive in the water. This long dive is called "sound".
After one " sound ", surface d~ves are repeated several time. The time'of " sound " is relaively constant, depending on kind of whale, in case of
baleen whale, it is from 7 to 10 minutes, but this exceeds 30 minutes incase it senses danger. It is especially long in case of sperm whale, which
is an tooth whale. It is from 30 to 70 minutes and averages about 40
minutes.
Several explanations are given on how oxygen is supplied during suchprolonged submergence.
OmmanneyCl) states that the vascular networks (retia mirabilia) functions as this oxygen supply. That is, a special structure called vascular
networks which are abundant in blood vessels and fats ac~ as accessory
lung and functions as strage for oxygen. However, Laurie(2) states that
this is impossible . and points out that from the relation between. basal
metabolism and lung's vital capacity, whale can be submerged for a longtime. That is he calculated the lung's vital capacity of a 122,000 kg blue
whale at not less than 3050 liters and that the air requirement per minuteat 178.75 liters. He states that according to these calculations, it is possible
for a whale to be submerged for at least 17 minutes.
As stated above, a whale makes several surface dives and then a longdive. "sound" From this fact, the author considers that the whale breathes
several times during the several surface dives, by this stores a large
quantity of oxygen in some part of the body and consumes it during a long
submer gence. One of the storage organ may be the vascular networks as
proposed by Ommanney out the author thinks that it is rather the respir-'

On the Respiratory Pigments of Whale97

atory pigments. The muscles of mammals which carry on a swiming life
such as pinnipedias and cetaceas are darker in color than those of landmammals. Especially in the case of sperm whale, the muscles are almost
black in color.
The so-called respiratory pigments are haemoglobin, myoglobin (muscle
haemoglobin) and various kinds of cytochromes. All of these are pigmentproteins containing iron and conbines readily with oxygen. Of the above,the molecular weight of myoglobine is 17,500(3) or about 1/4 of that of
haemoglobin, about the same as that of haemoglobin in iron content, or
0.345%,(4) combining power with oxygen and carbon monoxide is muchstronger than haemoglobin and also, reaction is much faster.(5) There areseveral kinds of cytochrotnes and it is believed that they act as carriersof hydrogen by· the oxidation and reduction of iron within the molecules.
The representative ones are a, b and· c.
The author carried on quantitative analysis of iron .in haemoglobin,
myoglobin and cytochromes a and c of these respiratory pigments anddetermined the content of the different pigments. As control, the bloodand muscle of cow and horse, which are land mammals, were used.
To determine the haemoglobin content in blood, Sahli's method using
haemoglobinometer was used concurrently with the method in which ironcontent is quantitatively determined directly. The result of the determination shows that by Sahli's method, there is no great difference inhaemoglobin content between whale and land mammals.
However, thedirect determination of iron indicated that it is considerably greater inwhale than in land mammals. In the case of land mammals the resultsfrom Sahli's method and iron determination method coincides rather closely
but in the case of whale, the iron content is greater in each case. That
· is, it .can be assumed that there is a greater quantity of iron compounds
other than in the haemoglobin state. By Barkan and Schales,(6) it was
discovered that in the blood. of ma,.mmals, desides haemoglobin, iron-containing organic compounds whose iron ion is readily liberated by dilutehydrochloric acid are also present. It is said that this substance has anaffinity to oxygen a,bout 4 to 10 times greater than that of haemoglobin.
It can be considered that perhaps the presence of these substances isanticipated.
In regards to cytochrome b, the quantity was so small that it was

98

Tadashi Tawara

impossible to determine so it will be omitted. Content in cytochrome aand c did not differ greatly from those of land mammals. Thus it 1s
assumed that it did not have any special· meaning relative to oxygenstorage.
Myoglobin content in whales is much greater in all cases when compared with land mammals and especially in the case of sperm whale which
ma.kes "sound", the content is about 8 times more.
From· the above result, it is assumed that one reason Why whales can~ithstand " sound" is that oxygen is stored by myoglobin within the muscles.
The author wishes to thin:k Dr. Tsutomu Naruyama and Mr. Tadashi
Nakai for guidance in this experiment and to Profs. Shichiro Akiya andTeizo Ogawa for their instructions. The author is grateful to Mr. Ryusuke
Fukazawa for assistance in the experiment and to the people of the
Ayukawa Station of the Taiyo Fishing Co., for assistance in providing
materials.
Experimental
Materials used in this experiment was a sperm whale and a sei whale
caught off shore of Kinkazan Island, the former 31 feet o and the latter
45 feet o . In both ·cases, they were 24 hours after death.*
Materials from cow and· horse used as control was obtained immediately
after slaughter from the Shibaura Slaughter House in the case of heart
and in the case of muscle, those on the market were used.
(1) Quantitative IX:ttirmination of Haemoglobin
Blood samples in each case was obtained from the heart.
(A) Sahli' s Method
According to Sahli's method, haemoglobin was transformed into acidhaematin by adding ~- HCl and colorimetric determination was madewith Sahli's Haemoglobinometer.
(B) Quantitative Determination of Blood Iron
In accordance with Pincussen's method,(7) blood was decomposed with·,sulphuric acid and 30% H202, and comparative colorimetric determination
was made of it, colored with KCNS, with standard concentration solutionof Iron Alum (FdS04)3 (NH4)2 S04•24RO) colored with KCNS, using* Materials were obtained in July, 1948. After this, there was a revision in law and cathing of
sperm wh&le les$ than 35 feet was prohibited,On the Respiratory Pigments of Whale99

Duboscq's colorimeter.
Table 1.

Sei Whale
Sperm Whale
Spem Whale foetus
Cow
Human(29 years old)

Quantity of Haemoglobin (in 100 cc of blood)
Haemoglobin

Fe (Hbx0.336)

Fe(Pincussen Method)

15.6g

52.4mg53.lmg19.Smg41.6mg43.7mg84.0mg95.0mg53;smg41.0mg46.5mg

i5.sg
5.8g
12.4g
13.0g

Quantity of haemoglobin determined by Sahli's method multiplied by
0.336 should give the quantity of iron in the haemoglobin state.
In the·case of cow and human the quantity of iron in the haemoglobin state andthe total iron content is about the same. However, in the case of whale,iron determined by Pincussen's method is much greater. The differenceis great especially in the case of foetus. It is believed that this is. becauseof the presence of larger quantity of iron other than in haemoglobin state.
(2) Quantitative Determination of Iron in Cytochrome State
In order to be aC"curate, the quantitative determination of cytochromeshould be made by spectral analysis but in the case, it was refined considerably and iron content determined. The determination figures· does notindicate absolute quantity but relative quantity determined under the same
conditions. For materials the above-mentioned heart and muscles of whale,
cow and horse were used.
(A) · Quantitat~ve Determination of Iron in Cytochrome a StateCS)
A solution containing 2% Na-cholat and -21Q- mol Na2HP04 is, forconvinience, designated as A-solution. Muscle is washed with water,
chopped with a meat chopper and 1 kg taken. 3 liter of A-solution isadded to this, extracted for one night at room temperature, strained withgauze and to 3 liter of the filtrate, 750 cc of a weakly ammoniac saturated
ammonium sulphate soltion is added (0.2 saturation). ·This liquid is· separated with a centrifugal separa!or and to the clear liquid is added
more saturated ammonium sulphate solution to make it 0.5 saturation.
Cytochrome a will precipitate out. The precipitate is placed in · a centrifugal separator, disrnlved in 200 cc of A-solution and made 0.2 saturation with saturated ammonium sulphate solution. This solution is left
standing for 2 days in an ice room, precipitate is removed with centrifugal separator, the clear liquid is mad~ 0.33 s;ituration with ammonium

Tadashi Tawara

100•sulphate, precipitate is dissolved with 100 cc of A-solution, precip:tateobtained by 0.2 saturat'on ammonium salphate is removed, precipitate
obtained by 0.33 saturation is collected and this operation is , repeatedonce more. Finally, precipitate obtained by 0.33 saturat:on becomes a
reddish brown, paste-like substance. This is dissolved in A-solufon, thetotal quantity brought to 100 cc., 10 cc. of this is taken, the entire precipitate obtained by saturation .with ammonium sulphate is decomposedwith HS04 and H202 and iron determined by the aforement'.oned Pincussen's method.
Table 2.

Quantitative Determination of Iron in Cytochrome a State (mg/kg)
Sei Whale

Sperm Whale

1.29
2.54

1.91
4.03

/Cow
··~-

Body muscle
Heart muscle

!

2.28
0.27

In the case of CO\y, the figure for body muscle was larger than forheart. In this case, it was not possible to obtain body muscle ar:d heartfrom the same cow so the materials came from different cow.
(B) Quantitative Determination of Iron in Cytochrome c State(9)(10)The· muscle sample is washed with water, chopped with a meat
chopper, 2.5% trichloracetic acid is added to 1 Kg of this, stirred andextracted for about 2 hours at room temperature. This is then strained.with gauze and 1 liter of the filtrate is brought to pH 7.0 with 6N-NaOH
solution. 500 g of ammonium sulphate is· added to this and left standingfor one night in an ice room. Thrs is centrifugally separated and theclear liquid brought to pH 3.7 by adding 5N-H2S04. Precip'tate preduced
from this is separated with centr;fuge, 20·cc. of water is added. to thisand· dialyzed (cellophane) against 1.% saline solut'on for 2 days. The
liquid. thus obtained is centlifugally separated and a dark red, clear
cytochrome c solution is obtained. This is diluted to 100 cc., 10 cc. of
this is saturated with ammonium sulphate at 60°C, the entire precipitate
is decomposed with H~so4 and 30% H202 solution and quantitative
determination of iron made by the aforementioned method.
Table 3.

Quantitative Determination of Iron Cytochrome c State (mg/Kg)

Sei Whale j Sperm Whale__B_o_d_y_m_u:C1-e---i--0-.4_3__
0.47

Blue Whale
0.44

iI

Horse
Cow
--0-.3-0_ _ ,._ _ _ __1_.i!__ ____H_e_a_rt_m_u_sc_Ie______
1._29_~~-o_.5_7_~--~·-~!_ _ 0.70 _____

On the Respiratory Pigments of Whale101

Cytochrome c is obtained by multiplying this quantity of iron by ~~4~ .
(3) Quantitative Dete:rmination of Iron. in Myoglobin. StateC4)
Sample muscle is washed with physiological saline solution, chopped
with a meat chopper and l kg of this is taken. - l liter of water is added
to this, extracted for l night at room temperature and strained with gauze.
N-NaOH solution is added to l liter of this solution and brought to pH 7.0,
250 cc of concentrated basic lead acetate solution is added and the precipitate produced from this is separated with centrifuge. Na2HP04 solution
is added to the clear liquid while maintaining it neutral with NaOH andlead is removed. 5 cc. of the clear liquid obtained by centJ:ifuge is taken,
saturated with ammonium sulphate, the entire precipitate is decomposedwith H2S01 and 30% H202, and colorimetric determination of iron, in theform of Rhodan-Fe, is made.
Table 4.

Body muscle
Heart muscle

Quantitative Determination of Iron in Myoglobin State (mg/kg)
Sei Whale

Sperm Whale

Cow

I

31.35
7:62

151.00
23.12

18.08

I

Horse

I

- - - - - - - - - - - - - - - - - - - - - - - - - - - -------------- ··----------

--

I

7.15
-

---- ----- - - - - - - -Myoglobin is obtained by multiplying this iron quantity by ~~ .
5Compared to cow, the myoglobin content is 2 times more in sei whale, and
8-9 times more in sperm whale.

0

References( 1) Ommahey F. D.: Discovery Rep. V. 329 (1932).
( 2) Laurie A. H.: Discovery Rep. VII. 365 (1934).
( 3) Theorell H.: Biochem Z. 268. 46 (1934), C. A. 28. 2736 (1934).
( 4 ) Hugo H., Theorell H. : Ibid, 252. 1 (1932), C. A. 26. 5587 (1932).
( 5) Theorell H.: Ibid, 252. 1 (1932), 268. 64 (1934), C. A. 26. 5587 (1932), 28. 2737 (1934).
( 6) Barkan G., Schares, 0.: Z. Physiol. Chem., 244. 81 (1936), 244. 257 .(1936), 248. 96 (1937), C.
A. 31. 715 (1937), 31. 1051 (1937), 31. 7898 (1937).
( 7) Pincussen: Mikromethodik, 108 (1930). Sudo, K.:lgakuteki Biryo Sokuteiho 155 (1937).

( 8) Yakushiji E., Okunuki K.: Proc. Imp. Acad. 17. 38 (1941)•.
( 9) Keilin D., Hartree E. F.: Proc. Roy. Soc. (London), B 122. 278 (1937), C. A. 31. 5390 (1937).
(10) The~rell H. : Biochem Z. 298. 260 (1938), C. A. 32, 969 (1938).

Research on Methionine in Whale(On Methionine (CH3S-CH2-CH2-CHNH2) Content)

I

COOH

Masami Yoshida

I. Introduction
Methionine is an amino acid containing sulphur and was discoveredin the hydrolyzed product of casein by Mueller in the United Statesin 1922. This .was synthetically produced by Bagar and Coyne in 1928
and named it methionine. After this, a large number of people attempted
to synthesize and isolate it.
From the nutrition point of vlew, methionine, together with cystine, isan ,important amino acid as a source of sulphur for animals. According
to recent researches, it is said that methionine transforms into cystine ina living body. In 1937, Rose(I) reported that methionine is a necessity andcystine is not· a necessity and explained that it aids growth, and disproved
Osborne an Mendel's theory.·
Furthermore, in regard to its content, Baernstein in 1932, made determinations of methionine, a characteristic methylthiol group, of various proteins
and reported that methionine content is greater in animal protein than invegetable protein.
. 1.8%(2)2.3%(3)Glycinin
Lact albumin
2.0%(2)
4.5%(3)
Gliadin
Egg albumin
2.2%(3)
3.1%(3)
Zein
Casein
3.14%(4)Sardine flesh
3.66%(2)
Beef
Whale meat (Kind unknown) 2.92%(4) ·In regard to methionine content in whale meat, the only report is by
Tomiyama.
The auther made determinations of methionine content in whale meatin order to determine its nutritional value, compared with other foodstuff
and as a first step in effective separation and utilization of methionine
from whale meat; a relatively detailed analysis· of its content in various
parts of sei. whale, sperm whale and fin whale was made.
There are various ·methods for making this determination, but Baernstein's method was used here.
\

Research on Methionine in whale

103

Baernstein made quantitative analysis under the assumption thatmethoxyl and methylimid bases does not exist in protein molecules andthat ·the quantity of methyl iodide obtained by reaction with HI results
only from CH2S.
For accuracy, methionine content in beef protein and casein wasdetermined and compared with previous literatures.
II. Experiment,.
(a) Collection of Samples
Heart, tongue, ventral meat (Sunoko) and ·meat was obtained from a
13.6 meter male sei whale at Ayukawa, Miyagi Prefecture. The length and sex of the fin whale and sperm whale are not known, but both wereobtained at Ayukawa. For beef, the round of beef obtained on themarket was used.
(b) Preparation of samples
Whale and beef protein
Flesh is fitst sliced thinly and soaked in cold water, wh'ch is heatedto 36°C and changed to new water. The it is bo]ed once. This is thenexpressed and water removed as much as possible, I chopped in a meat
chopper, dried in sun, extracted with boiling alcohol for 4 hours andthen with ether for 10 hours and ·dried for 3 'hours at 105-110°C.
Balem
After washing with water, it is dried at room temperature, cut in thinslices, extracted with ether for 10 hours and dried for 3 hours at 105ll 00C.
(c) Outline of Operation
Sample is decomposed with HI and CHJ produced, absorbed in alcoholic AgNO:i solution to form AgI, filtered and the residual Ag determined
quantitatively with 2/100 N KCNS using m:cro-burette, The methionine
quantity is calculated from the difference. in Ag· quantity in a blank test.
The cond:tions for the determination were a.s follows ; In the originalreport, it is stated that about 90% of the CH:I come over in the 1 sthour but the remainder usually takes several hours more, and Baernstein
adopt about 15 hours, but here the reaction time is 6 hours, about 0.5 gof the sample is decomposed with 10 cc. of HI (s. g. 1.7) and absorbed
in 20 cc. of alcoholic AgNOs solution.

Masami Yoshida104

The apparatus is as follows :I

I

AgN03 Solution

20% CdS0.1 Alcoholic AgN03 Solution

H2S01(d) Experimental results
Table No. 1.
Content

SampleHeart
TongueVentral meat (Sunoko)
Belly meat
Head meat
Back meat
Tail meat

Sei Whale

4.80
4.22
4.17
3.42
3.33
2.78
2.71
4.82

Fin Whale Meat
Sperm Whale MeatBlue Whale Bajeen
Fin Whale Baleen
Sei Whale Baleen

3.46
0.82
1.92
1.96

The methionine content of beef and casein was determined by thesame method and compared with previous literature, it "is as follow.s :Table No. 2.,
AnalystSampleCaseinBeef protein
Whale meat protein

Baernstein<2)

Baernstein(a)

3.53%
3.36
3.25

3.1%

Tomiyama,
HanadaC4)

Yoshida

2.54%

3.94%

2.92

3.06

3.66

3.56

In Table No. l, the average methionine content of 3.06% in the various
parts o_f s~i whale, which is ordinaray called meat is very close to. thevalue 2.92% by Tomiyama, · which was obtained by the difference between

Research on Methionine in Whale

105

total S for Denis Benedicts method and cystine S fer Okuda's method.
However, there is something here which require special attention. In
the case of casein. several experiments' gave the result of 3.94±0.02%, but
in the case of beef and whale meat, there are some cases when the resultsof several experiments does not coincide. As to the cause of this discrepancy, on considering Table No. 1, it appears that there is a tendency forthe value to be h;gh in parts '\Vhich are rich in connective tissues. Thus
it was thought that there may b.e some relation between ·methionine content and the connective tissue contained in the sample, so the following
experiment was performed.
The kind of whale is not known but the sample was prepqred in thesame manner as before, connective tissue was separated as much as possible from the meat and methionine of each was determined. The result
is as in Table No. 3.
Table No. 3.
Connective Tissue

Meat Scrap

3.22%

1.43%

That is, from. the determination results, it is clear that methionine
content is greater in the conective tissues than in the meat scraps.
III. Summary(1) The methio.1ine content of the part ordinarily called meat is 3.06%
in sei whale, 4.82% in fin whale and 3.46% in sperm whale.
(2) Of the different parts of sei whale, content is largest in heart, next
in the portion rich in connective tissues (tongue, ventral) and then in
the part where there is little connect~ve tissues (meat).
(3) The difference in methionine content in the different part of whale
meat is thought due to the difference in the quantity of connective
tissues in the various parts.
(4) The large content in the heart is thought to be of great interest in
relation to physiological action of renewal.
Refferences
( 1)
( 2)
( 3)
( 4)

Womack Kemmerer: J. Biol Chem. 121 (1937) 403.
Baerstein: ]. Biol. Chem. 97 (1932) 663.
Baerstein: ]. Biol. Chem. 115 (1936) 25.
Tomiyama Ha~ada; Nogei Kogakukai Shi 10 (1934) 53.

Factory Ship Whaling Around Bonin Islands m 1948
Kazuhiro Mizue
1. Introduction
.

.

.

;

The ·author carried on investigation on board the whaling factory ship
Kaiko Maru of the Nippon Marine Products Co., which carried on whaling
in the waters around Bonin Islands from February to May, 1948.
Various bfological investigations were made on bo~d the factory ship
and a general outline will be presented here.
Catcher boats attached to the factory ship were the No. 2 Kyo Maru
and No. 3 Kyo Maru of the Nippon Marine Products Co., and those attached to No. 9 ship of the Taiyo Fishery Co., were the Seki Maru, No. 7
Seki Maru, and the No. 7 Seki Marn was replaced by the No. 2 Seki Maru
during the operation.
Salting and refrigeration ship was attached to both fleet to act astransporting ships, and in general, the composition of the fleet was asabove.
First, a brief history of whaling in the waters around Bonin Islands isa follows. Whaling was begun in Japan__ before the preceding century but
whaling in the waters of Bonin Island was of relatively recent origin. In
1922, ·an investigation was made ~o determine latent resources in the watersof Bonin Islands as a whaling ground, and actual operation was begun in
the following year, 1923. That is, the Toyo Whaling Co., the predecessorof the present Nippon Marine Products Co., was the first to develop thewhale resources in this region. Much later, in 1937. Taiyo Fishery Co.,
known as Hayashi Kane Shoten at that time, also engaged in this operation.
From then, the so-called Bonin Islands base .whaling was continued
until 1944. During this period, Nippon Marine Products Co. maintained a. base on Chichi Jima and Taiyo Fishery on Haha Jima, competition between
the two was removed and operation was carried on in a friendly manner.
At the time when land bases were established and operation begun in
1922, it ·was a good fishing ground for winter, shore approaching humpback whale, similar to those offshore Galanpi Cape in Formosa, and together
with this, the whaling ground was developed year after year, so that catches

Factory Ship Whaling Around Bonin Islands in 1948

107

of Sperm Whale and Sei Whale gradually increased and considerably goodwhaling record was established, such that after 1939, a total of 200 whales
were caught.

.

Table 1.
1922-1946 Monthiy Wahle Catches for the Bonin Islands
Season1922-23
19:i3-24
1924-25
1925-26
1926-27
1927-28
1928-29
1929-30
1930-31
1931-32
1932-·-33
1933-34
1934-35
1935-36. 1936-37
1937-38
1938-39
1939-40
1940-41
1941-42
1942-43
1943-44
1944-45
1945-46
Total

November December

0
0
0
0
0
0
2
0
0
0
0
0
1
0
0
0
0
0
0
'()

0
0
0
0
3

January

Februaly

March

April

3
3
0
0
0
5
0
14
1
5
1

0
9
21
12
5
8
6
18
16
16
26

0
29
17
19

0
28
28
20
9

0
17
23
12
2

4
8

4

11
13
5
17
1
3
0
0
0
0
0
0
0
82

10
10
11

18
23
44
9
7

20
19
11
0
0
319

5
9
6
3
12
13
25
16
16
22
36
11
67
40
72
37
36
32
0

5
11
11
16
9
23
28
42
16

0
0
0
0
0
4
1
25
18

22

55

34

141

50
93
0
82
67
0

52
6

115
158
0

!

May

0
8
6

7
0
0
0
0
0
0
0
0
0

2
0
6
8
4
5

0
18
0
0

I ·Total
3
94
95
70

21
30.
22
40
40
45
68
50
64
93
131
79
211

244
229
63
270
268
0

Table 1 shows the yearly catches of whale by month, from 1922 when
operation was begun, up to 1948.
Jn regards to Table 2, only one company was engaged in whaling fromthe time whaling was begun, up to 1938 so that yearly average number ofwhales caught was only 62. However, after Taiyo Fishery began actual
operation in the following year, the rate of catches increased :i;apidly forthe following three years and the yearly average catches increased to 228
whales. The following year was the year in which war began and withthe outbreak of war, operations in the waters were greatly' restricted. As
the result of this, the catches shows a sharp decline and only 68 whales
were caught during this year. Operations were again resumed to normal

Kazuhiro Mizue

1-08

Fig.
1
Whale Catches Around the Bonin Islands, 1922-1948,_ _ _ _ .Q ~itpC:n_M~r~n"- ~~c~ E_o~ _ _ _ _ _ _ _ _ _ _ _ _ _ _00\'22Zi! T~iyo Fishery Oo.

- - - - - - - - - 200'- -

-

-

-

-

-

-

.

-

-

-

- - -

-

-

-

-

-

-

-

--

·-

-

-

-

.

-~

,

-

-

,~--f-.
t-.
" -

-~ -

.c

j
0

-~

-

I
-----1---- - - - - - - - - - - - - - - - - - - - - -~ - - -1- ·100

-

-

-

.;
z

l

-

-,

"'"'
"'
°'..,"'

-

-

-

-

-

-

-

-

-

-

-·~ --1------ - -

nnnnnn"' "'"' ... "'

-

7

j

-~-

-~-

I

; -- ~ - lI

-

~

I
/
'/
Y.
,_
-T
....
0.,..
!(;' ~"' "' "' "' "'"' "' ~~7~7~~:~~;"' "'"'I "' "'
"'I
,_
...
... "'' "'
....
0
"'
"'
"'
"'
"'
"'
"'
"' ~"' ~"' ~ ~"' ~"' ~"' ~"' ~"' ~ ~"' ~ ~ ~"' ~~~~;g~~
~

,_

I

l

I

I

I

I

I

i

I

the following year and the year after, inspite of being in the midst of war
and with the increase demand for the army and domesticuse, a record
number of catches were made.
The following year was the year in wh~ch war was terminated and base .
whaling in the Bonin Islands were stopped with the fall of Saipan andwith the occupation of Iwo Jima by the United States Forces. During thisperiod, (22 years), the average yearly catches were 155 whales .
.I~ J'.~~,'!ls to the whaling season, as can be seen in Table 1, the total
catches is greatest in March, followed by February and April.
As to kinds of whale, ordinarily, they are Humpback Whale, Sei Whale,
and Sperm whale and sometimes, Blue whale and Fin whale are caught.
In regards to whaling ground, the center of operation is 100 nautical
miles east· of Ha:ha Jima. (It is needless, to say that Humpback Whales
have the habit of approaching the shore so this is an exception).
\With the termination of war, the Bonin Islands wer~ outside the area
in which fishing is permitted. However, due to the zeal of the whaling
firms and efforts of the Fishery Bureau,, and due to the kind consi.derationJ'Factory Ship Whaling Around Bonin Islands in 1948

109

of SCAP, factory ship whaling in the waters of Bonin Islands was permittedwith various conditions attached. The result of whaling after the use offactory ship whaling is as shown in Table 2.
Table

2.

Fin Whale ( Se1· W h ale~ -Blue-Whale
--

y

1946
1947
1948

!

4

Humpback. Whalei

29
150I•SprmWhale !_ Total
I---------

I

12
1
3

105

3

1

I

12
195
1 31

113
350
243

As mentioned above, many Sperm whales, Sei whales and Humpback
whales are found in this fishing ground and Blue whale, Fin whale andRight whales only rarely migrate here. After the termination of war, operation was. changed to factory ship 'whaling and the number of catches of
Humpback whales was greatly reduced, as compared with the period when
whaling was by base whaling. This is a natural result because Humpback
whales likes to migrate close to the reef of the island and since factory
ship '''haling cannot be carried on within 12 nautical miles from shore.
During the base whaling period, operation was carried on from NoveFig.
2a
Lbcation of Catch of Sperm Whale
El42

El44

El45

~CHIC HJJIMA.~!\r HAHA
0~· JIMA

.

.

.

I

..

N27

'11,

a

• "•t'I>
8.. et· .

..

~

!!I

.
...

·-

..

·;

:
l\

;

..

IF
!I

N26

......March•····April

N25

110

Kazuhiro Mizue
Fig.
2b
Location of Catch of Sei Whale
El42

El44

.•• --March.0--.April~~~-+~~~~~~t--~~~~~-t-~~~--~--r~~~~N~mber to May of the following year, the peak being from February to April.
Even with the change to Factory ship whaling, this peak period is taken
advantage of.
Oiiginally, the fishing ground in the Bonin Islands is very small,'compared to that off shore Sanriku Area. Table 4 is a chart showing theposition of catches in 1948 and in looking at this, catches were madebetween lat. 26cN-27°N, long. 143"E-144°E, or in an area of 60 nautical
miles squaee.
That is the principal whaling ground is 50-100 nautical miles E-SEof Haha Jima.
Also, in looking at this chart, the position of catches of Sei whale
and Sperm whale gradually approach the islands from February to May.
This is related with the water temperature and feed.
The quantity of fuel oil was very small in 1984 so during the 1948
whaling, the factory ship could not be stationed in the middle of the fishingground and usually was in a drifting condition so in looking at the positionof both factory ships in the course chart, it is apparent that they driftedFactory Ship Whaling Around Bonin Islands in 1948

111

here and there at the mercy of the wind. As a result, both fleets were
cosiderably separated from each other and resulted in numerous difficulties
in operation.
2. DetailIn 1948, besides Sperm whales and Sei whales, only 3 Fin whales.
1 Blue whale, and 3 Humpback whales were caught. This is a very small
number, and in the future, it will be necessary to investigate, compare andexamine it with those of Antarctic Ocean and investigate it from theresource point of vlew.
(a) Sei whale-Balaenoptera borealis (Lesson)-Table 3 shows the length of whales caught and of the total of 105
caught by both fleet, sex was about half and half.
Table
Body length in feet
35
36
37
38
39
40
41
42
43
44
. 45
46
47
48
49
50
51
Total
Mean length

3.

0

2
2
1
3
2
7

Total

l

1
5
3
6
6
2
16
19
10
10
8

3
1
5
3
9
9
3
4 .

10

7
6
3
5
3
3

54
42.65

~

i

I

5
6

11

1

3
3
1

1
51
41.96

1
105
42.31

Mean length for 0 was 42. 7 feet and 9 w.as 42.0 feet,
Mean length for 0 and 9 as 42.3 feet.
The largest whale was 48 feet for 0 and 51 feet for 9 and the smallest whale was 36 feet for o and 35 feet for 9. The peak of body length
frequency curve is 42 feet.
In such Sei whales, there is a considerable difference in length with

112

Kazuhiro Mizue

sea areas, and average length is greater as it goes northward and getssmaller .as it goes southward from Kurile Islands sea area, Sanriku sea
area, Bonin Island area, Kinan sea area and Tsushima-Formosa sea area.
In viewing former statistics, there is as much difference as 5 feet in tnose
from north s.ea area and Tsushima-Formosa sea area. The average lengthof Sei whale caught in the .waters around Bonin Islands in 1948 was in
between these.
Also, in Sei whales caught in the waters around Japan there are somein which corrugated markings on the dorsal and ventral side are conspicuous and some which are not. Thoese which have corrugated markings
are called "totan-iwashi" (corrugated Sei whale) and as a general rule,these are of Northern variety· and those without corrugated markings· arefound in the south western sea area and are of southern variety. It isthought that northern and southern varieties may be differentiated fromthese corrugated markings and difference in length.
However, during the current investigation, Sei whale with corrugatedmarkings are ·not limited only to large whales and considering other points,
there are no indications that theY. are of different speice. That .is, theso-called corrugated Sei whale are Sei whales with numberous old scars
and are not regional difference or sub-specie. Scars wh:ch are the causeof this corrugated markings are due to parasitic protozoa and in extreme
cases, there are considerable number of scars so that they can be scooped
up with a spoon.
Outside of this, the only exterior parasites were small number of pennella whose part exposed from the skin surface, has fallen off. As ageneral rule, exterior parasite is very few because of sub-tropic area.
Furthermore, as to interior parasite, only a small number of distoma
was discovered in the small intestine.
In regards to these parasites, it is planned to report on this after
research, together with those in the stomack.
As to thickness of blubber, the vertical thickness was measured for
99 whales at the s:de below the dorsal fin and back of the head. The
mean value below the dorsal fin was 5.2 cm and are of normal nutritional·condition in general. In regards to investigation of the mammary gland,
it will be omitted here.
As to feed, there were ·many with empty stomach and for 140 whales

Factory Ship Whaling Around Bonin Islands in 1948

113

investigated, 40 had empty stomachs, 13 had eaten Euphausia, 34 Calanus
and 17 young sardines. As a ·general rule, kind of . feed was of single
variety.
As to the investigation of reprodduction gland, the weight and volumeof the o testes was determined. Table 4 shows the relation between bodylength and weight of testes.
Fig.

3

Relation beween the weight of the total of rightand left testes and body length of Sei whale.

..

•oooo

.: .
. . .. .
..

600
40oo·
2000
gr.

t

• l • • .•
I

;15

40

~Body·length in feet

That is, it is about 1 kg (total of left and right) up to 39 feet andmore than 4.0 kg above 42 feet. In general, it is said that the sexual
maturity of o Sei whale is 44.6 (13.5 m) but from this investigation, it isestimated to ·be 42.0 feet. Howeber, there is a very marked difference in
the Sei whale in different regions so it is necessary to make comparative
studies with those of other regions.
As so the 9 , the weight, volume, number of corpora and graafianfollicle of the ovary were determined.
According to the corpora Table showing the relation between corpora
number and length of body, the body length of minimum sexual maturityof a 9 is 38 feet, minimum for pregnant whale is 41 feet and maximum
is 46 feet. Even when looking at the frequency curve of corpora number,the number of young whales with little corpora number in very large asa general rule.
As to the weight of the ovary, there is considera~ledifference in accordance with the presence or absence of functional corpora
but generally they are matured at 250 gr. It is said that as a general rule,the body length of sexual matqrity of a 9 Sei whale is 14.5 m (47.9 feet)but according to the present investigation, it is much smaller than this andis· completely matured above 46 feet. Also, in case there were. those withfunctional corpora the uterus was always out open, inspected for foetus andKazuhiro Mizue

114

their. body length, sex, etc, were· determined. Pregnant whales numbered11, smallest foetus 2 inches and the maximum 3 feet 5 inches. Their sex
were 6 o , 5 9 and no multiple foetus or deformity were noticed.
Next, the adhesion of the cartilagenous layer between the vertebral
matter and tip of bone of the vertebrae was determined and the maximum
body length of non-adhesion in .o was 46 feet and the minimum bodylength of adhesion was 42 feet. In the· 9 , it was 49 feet in the former
and 46 feet in the latter. In the o, there were some which had adhesion
at 42 feet.
(b) Sperm Whale (Physeter macrocephalus, L)
Sperm whale is polygamous and has a habit of forming harems. They
are distributed. widely throughout the world and in the waters around
Japan, from offshore Sanriku to Hokkaido and Kurile Islands are goodfishing grounds. It is the general rule in all myiiitacoceti that 9 is larger
than o, but in the odontoceti, this is the opposite. In Sperm whales, ois much larger than 9 .
Past statistics shows that large sperms of more than 60 feet are notrare buf maximum for 9__ is 31 feet and there are only a few above 40 feet.
Therefore, the body length and production of sperm whales-differ cons:derably with sex so that they should ·be considered as of being of different
variety.
Whalers in Japan classify Sperm whales into the t.hree classes of large
sperm, medium size sperm and small sperm. Table 4 shows the length of
Sperm whales.
Fig.

4

Body length in feet

~

Total

26
27
28
29
30

1
1
1
2
3
6

31

1

32

l

33

2

1
1
1
1
3
5
6
6

34
35

4

27

36

3

37

l

13
8

1

3

.•

7
8
3
3116
9

Factory Ship Whaling.Around Bonin Islands in 1948

115

Body length in feet

0

fj!

Total

38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
Total
Mean length

1
1
3
1
1
1

3
3

4
4
3

1
2
3
2
3
2
3
4
6
I
1
1
50
44.64

i
1
1
1
2
3
2
3
2
3
4
6
1
1

!

1
81
34.41

38.31

The total catches for both fleet was 131 and of this, 50 were 0 and
81 were 9 . The size of the Sperm whale harem in the vicinity of Bonin
Islands is much smaller than those off shore Sanriku, but even then, it is
not a rare thing to see schools of 20-40 small Sperm whales. However,since the catching of whales less than 35 feet by factory ship whaling is
prohibited, the catching of 9 is restricted to a considerable extent andwhaling is done for o .
Mean body length is 44.6 feet for o and 34.4 feet for 9 .
Maximum body length for o is 57 feet and 39 feet, for 9 .
Minimum body length is 29 feet for o and 26 feet for 9 .
In picking out what appears to be the peaks of body length frequency
curve, it is 35 feet, 40 feet and 53 feet for o and 35 feet for 9 , but
determination of age school is very difficult due to insufficient information,
the same as in the case of Sei whale. In regards to color of body, Mr.
Omura is investigating and classif~ing Antarctic sperm whales but in thecase of sperm whale in Bonin Islands it differs with whales, and· various
kinds of scattered patterns and white spots are present around the stomach

116

Kazuhiro Mizue.

and pudenda. Howtver, these patterns are found not only in the large oof th.e so-called isolated Sperm whales but even more in small Sperms.
Also these differences in pattern is also recognized from the time of foetus.
Next, Sperm whales have teeth and there are two 'rows, ih the lower
jaw but in ~he case of whales with short body length, these are buried
under the skin. Even in considerably matured whales, the innermost 2-3
pairs are not exposed but they appear only as protuberance of the skinwhen seen from the outside. The upper jaw has holes in which the teethof the lower jaw fits· in but 5-6 layers of traces of teeth can be recognized in it or in its v2cinity.
In many cases, the number of teeth on the left a~d right side differs ·but as a general rul~, there are from 12-25 pairs.
Scars of Sperm whale does not differ very much from that of Sei whale
but only in that the number is relat~vely small. Also, they are present
only in the tail portion in case of Sei whale, but in the case of Spermwhale some can be seen at the head.
Furthermore, in most cases, traces of line-like scratches can be seenaround the mough, which tells of fierce battles with large cuttlefishs, which
are its feed and coin-size traces on the suckers.
In regards to parasites the only thing is a few traces of pennella
parasite and 2-3 cases of conchoderma parasites of tooth. In case ofwhales from northern parts such as White. whale (not Delphinaptenes leucus
but old bull) Aosa whale (refers to North Pacific Fin whale with diatom
film attached), etc, forms diatom film on. the body and diatmes is greaterin whales in the lowen temperature belt and are parasitric and this is one
way of estimating ' the .period of life in low temperature .belts. From this,it can be said that whale in the Bonin Islands has very little relation with
the northern low temperature belt. As a general rule, there are very few
parasites on Sperm whale in this region because of its subtr~mic water,
similar to Sei whale. Also as to internal parasites, only the very common
ones are found.
Thickness of blubber was determined by the same method as in thecase of Sei whale and it was found that nutrition was ordinary.
Feed is ordinarily cuttlefish and oct-0pus but besides these,. there weresome which ate jellyfish. Those with empty stomach were 47 out of the
131 investigated but large amount of mouth piece and eye of cuttlefish andFactory Ship Whaling Around Bonin Islands in 1948

117

Nematoda were found in the stomach.
In regards to reproductive organ, the weight and volume of testes,
weight, volume, Graafian follicle corpora, etc were determined, the sameas in the case of Sei whale.
The testes of o Sperm whale is considerably larger than that of other
Mystacoceti, spherical in shape and its position is entirely different fromthose of Mystacoceti.
Table 4 shows the weight of testes and body length and the curve
makes a sharp rise from body length of 40 feet, and there are some wihch
exceed 15 kg.
Fig. 4
Relation between Weigit of testes (totalof left and right) and Body length of
Sperm whale

Fig. 5
Relation between Volume of testes (totalof left and right) and Body length of .
Sperm whale
~0000

15000

..

10000

5000
gr

Lb

. :.

...

20000

..

!0000

U,, . ~ ·~· . . , ,~.,,0~5404550
Body length ln feet

'55

60

. i .

Bod)

engtb 11' reet0

The same can be said for Table 9 also which shows the relation between volume of testes and body length.
There is considerable difference for each whale, but from th'.s, it canbe estimated that the sexual maturity of ~ Sperm whale is body lengthfrom 42 to 44 feet, as a general rule. However, more investigation isnecessary in order to arrive at a definite conclusiOn.
As to 9, the. body length of minimum sexual maturity is 28 ·feet,according to the corpora table. However, they are unmatured whales of
35 feet. Since the whaling was by the factory ship method and as aresult, catching was limited to whale whose body length is more than 35 ·feet, it is difficult to estimate the sexually matured body length by thisinvestigation alone because in 9 of Sperm whale, many· reach .sexual
maturity below 30 feet. As to corpora, those with 5 were most numerous,
centered around body length of 35 feet and maximum number is .14.

118

Kazuhiro Mizue

There were 22 pregnant whales, the maximum pregnant whale being
39-feet and 'the minimum 28 feet. Maximum body length of foetus was
13 feet and the minimum was 2 inches. Ratio by sex was 10 o .and 12 (f •It is reported that the length of pregnancy of Sperm whale is 16
months and can be roughly classified into the two groups of those whichwere pregnant for less than 1 year with foetus of less than 1 foot andwhose which were pregnant for a long time and approaching delivery, with
foetus of more than 10 feet.
However, there is a necessity of collecting more data to find out thegrowing period, accurate period, of pregnancy and body lenhth of foetussat delivery.
This ends the current report. The author wishes to express sincere
grasisude to Mr. Omura and Sakiura of the Fishery Board for direct
instructions in this investigation.

Statistic Study of Foetuses of \iVhales
Kazuhiro Mizue and Hisako Jimbo
Introduction
The material of the present study regarding foetuses of whales is based
upon the monthly reports forwarded to the Whaling Society by various
whaling companies from 1911 to 1948. Most of such reports were destroyed by the fire during the War. Thanks to Mr. Ohmura's efforts,
however, all the numbers issued in the following sixteen years, -1911,
1914, 1919, 1921, 1922, 1926, 1932, 1934, 1941, 1942, 1943, 1944, 1945, 1946,
1947 and 1948- have been recollected so as to be available for us atpresent. Moreover, six numbers of both 1910 and 1940, too, have been
restored.
These whaling reports record particulars of individual animales of themonthly catch with such items as date of the catch, species, sex, bodylength, foetus, stomach contents, and locality of the catch.
As for the older records of foetuses, they seem to have been madeaccording t? the rough. estimation by the eye. Yet there may be no grossforgery about them, as in the case of body-lengths which are subject. tobody-length limitation and ratio money. Since 1946, however, the records
have become authentic, because in that year whaling inspectors began tobe employed in each landstation along our coasts, who, as in factory-ships,
make biological investigation of the catch as well as superintend whaling
operations. As anyone that has been on board a factory ship and atpresent in the scene of dissection of a carcass, can well suppose, foetueses.
below 5 inches are very apt to be overlooked. In fact, either for this.
reason or for the ship in recording, description of such small foetuses are
extremely rare before 1945. Neither is it probable that before 1945 eachovaries with functional corpora lutea should have been operated, upon soas to ascertain the presence of a foetus. The descriptiop. of foetuses are
confined to such large sized species as Sperm, Sei, Fin, Blue, Hump-back,Right and Grey whales. Of these the foetuses of Blue, Hump-back, Rightand Grey whales are s.o few in number that it is hardly possible to study
them with any statistic result.

Statistic Study of Foetuses of Whales

120

Sex Ratio
As for Sperm whales, the ratio of males and females are 46.2% and53.8% respectively : so it is roughly half and half. Since early days it
has been said of Sperm whales, which have the habit of forming Harem,that females greatly outnumber males. · This is by no means correct.
According to the sex ratio of foetus of the same species, females slightly
outnumbe.r males, but this relation is quite reversed in the sex ratio of thecaught whale. (male:-i are 52.4%, females are 47.6%)
The sex ratio of the foetuses of Sei whales is 45.5% and 54.5%, showing · the ratio of either sex is almost the same. This relation is notchanged in the sex ratio of the caught whale. (males are 49.0%, females
51.0%)
Table 1.
Species

Foetus of Fin

Foetus of Sei

Foetus of Sperm

Year

Total
Male Female Unknown

T t 1
Male Female Unkna
own

I!Unkn- Total
Male !Female i
n

1910
1911
1914
1919
1921
1922
1926
1.932
1934
1940
1941
'1942
1943
1944
,1945
1946
1947
1948
T·>tal

Sex
ratio

----

0
1
2
0
0
0
0
4
2
4
6
6
4
3
1
1
1
3
38
51.4

0
0
2
0
0
0
0
1
4
2
2
8
6
2
1
6
2
0
. 36
48.6

·O
0
2
0
0
0
0
5
0
0
0
0
0
0
0
0
0
0
7

0
1
6
0
0
0
0
10
6
6
8
14
10
5
2
7
3
3
81
100

- - -°- - - - - ·' -ow

1
2

l

1
1

l

0

l

l

2
1
5
13
1
5
6
6

0
2
9
14

2

1
9
14
6
7
3
4

8
0

4
7
17
81
45.5

i
'

12
11
97
54.5

2
0
1
1
5
3
11
13
1
2
3
8
4
2
7
0
2
6
0
3
0
2
16
0
1
20
28
0
2
8
0
14
28
20
0
2
12
17
0
16
1
19
4
0
3
0
8
22
0
45
19
'
1 II 2956
I
21 I 199
247
46.2
100
!
I

I

5
3
7
4
9
9
3
2
25
13
38
4
14
21
9
29
33
60

!

'

I

I
I
I

0
1
10
0
0
3
0
3·

21
66
6
31
42

0
0

I

l

2
22

288
i
53.8'

18
3
5
45

0
0
0
0
0
2

i

I

6
7
18
12
16

I

13
51
79
118
557
100

-

So is the sex ratio of Fin whales, males being 51.4%. while females
48.6%. According to Mr. Matsuura, female foetuses are less in number
than male ones in Mystacoceti. But it cannot possibly be correct, for boththe sex ratio of foetuses shown in ·Table 1 and that of the caught whale,indicate either in Mystacoceti or. in Odontbceti the sex ratio is half andKazuhiro Mizue and Hisako Jimbo

121

half.
Relation between Pregnancy and Body-length
Fig. 1 shows graphi~ally the numbers of pregnant whales of the speciesof Sperm, Sei and Fin whales, respectively, according to their body-lengths.
For lack of material, we had to omit the observation of pregnancy in otherspecies. As for Fin whales, the minimum size of pregnant whales is 50
feet and the maximum size 71 feet, while the greatest number of pregnant
cases, are seen in the body-lengths of 64-66 feet. In Sei whales, theminimum size is 38 feet, the maximum size 56 feet and the peak of pregnancy is formed between 45-50 feet. In Sperm whales, the maximum size,of pregnant animales is 46 feet but females measuring 30-37 feet, especially
32-35 feet ·are most frequently pregnant. In the adjacent waters of Japanthe catch of Sperm whales below 30 feet is prohibited by body-length
limitation, while Sperm whales measuring 30 feet are ·already sexuallymature and therefore liable to pregnancy. For this reason the minimum
size of pregnant of Sperm whales cannot be determined.
In recent years the size of the whales in the catch has been reduced
Fig.
100

.1

~

llO

/,'\
'\
/,

70

.~ ! \

90- - - - - S'Pent ..'halo- - - - - - - - - - Sei Whsh

- - - - F i n \'hale

I' I \

'

60

5oi
40!
}OI
l

\' '

I \!i \I'\\

Statistic Study of Fo~tuses of Whales

122

as js clearly seen in any body-length frequency curve. So in the stock ofwhales as they are in the sea, this curve of pregnancy ~ay shift a little
to the side of larger sizes.
Growth Curves
Now we are going to determi~e the pairing period, the duration ofpregnancy, the delivery period and the body-length at delivery according
to such species as Fin whales, Sei whales and Sperm whales.
(A) Regarding ·Fin whales, Fig. 2 shows coordinate with the transversal
axis indicating months and the vertical axis indicating body-lengths offoetuses. The black dots representing the foetuses in our material, areplaced on the coordinates according to their size and the time of the catch.
Each signs of. CD represents an average body-length per ten days. Due tothe scantiness of our material, tracing these signs of CD would produce too
rugged a curve. So we have shown an average body-length per month bysmall circles, and prepared a growth curve based on them. We have no
foetus in our material for the month of November and of December. So
Fig.

2

..
10

'

·-

. .. . .

.. .
'

h••

I~monthf Ja;.""""~h~b.-'--:-:---'-A-~-.-'-~-,-1..~J-un-.-'--,J~ul-.J....-,A-ug-.'---,..-p.-'-~~--'-~No--'-~~.-. .i. . . ,.~~.
••

••

•

••

"

Kazuhiro Mizue and Hisako Jimbo

123

we are obliged to prolong the growth curve which has. already shown aclear tendency by the time it reaches the month of July-so as to complete
the year's curve. If we are to apply the body-length of Fin whales atdelivery, in the South Semisphere, 21-22 feet, determined by Mackintosh& Wheeler and Matthews, to our growth curve, the size will find its placeon the curve at the beginning of January next year. The pairing period
according to our growth curve comes in January. Therefore, if we areto assume,oand we have some reason to believe,-that the boby-lengths atdelivery in both the semispheres are the same, the duration of pregnancy is
twelve months and one decade. There is six months' variation in both theperiods of pairing· and of delivery between the South and the North Semispheres. Our growth curve has roughly two months' breadth.
· (B) The coordinates in Fig. 3 have been prepared in the same way as
those of Fin whales, and show the growth of foetuses of Sei whales. The
signs of CD represent average body-lengths per ten days, and the circles
those per month. The records of foetuses in the months of March, April
and May are exclusively those of the Bonin Islands sea area because there
is no foetus found in other sea areas duripg that period. The season for
Fig.

3

124

Statistic Study of Foetuses of Whales

Sei whales in the Bonin Islands sea area lasts only these three months andthe foetuses found then ,are still very small in size as is shown in .thecurve. In July, August and September pregnant whales are captured mainlyin the Sanriku sea area, and in October, mainly in the Hokkaido Padic
sea area. These shifts in seasons and whaling grounds as well as thedifferent growth stages of foetuses of the catch lead us to suppose that theSei whales in our adjacent waters migrate according to the course andseason above mentioned. If we had whaling landstation at the south of theBonin Islands sea area we should get still amaller foetuses.
During the whaling operations in this season, some marking weredischarged in the Bonin Islands sea area. The result of this experiments
will clarify the course and the season of migration in several years.
The growth curve of Sei whales was shown in Fig. 3, run smooth
upwards as far as the tenth month, after that the dots are hardly traceable .
. So we have prolonged the curve already drawn as in the case of Fin
whales. According to this curve, foetuses measuring 13 feet are seven in
number while those of 14 feet and ot 15 feet number one and two respectively. From this we assume foetuses at delivery measure 14 to 15 feet aridthat delivery time is at the end, of November. Like Fin whales, Sei whales
have their pairing period in January, so that the duration of pregnancy isless .than 11 months. Our growth curve of Sei whales has three month~'breadth.
Sei whales are to be subdiv:ded into totan whales (there are many
designs like the surface. of galvanized iron plate in abdomer.) and Sei whales
proper. Totan whales are larger sized and supposed to belong to a Northern stock while Sei whales are of a Southern stock and mostly captured
off the coast of Wakayama Prefecture. The ct:fferences of these two kinds
of Sei whale have been a subject of many d:scussions but not yet put toany morpholog~cal investigation. Fig. 3 show four dots placed far apartfrom the main .group. These four foetuses are all found in animals captured off Wakayama Prefecture. This will give rise to the question whether
the Sei whales in that neighbouring sea are of different species from thatof Totan '*hales or of the same species taking different course in theirmigration. The problem will be clarified by the further study of the localityof the catch, of the d:stribtion of body-lengths according to the sea areas"and 'by experiments 'by marking as well as by morpliolog:cal investigations.

Kazuhiro Mizue and Hisako Jimbo

125(C) In order to ascertain the courses of migration of Sperm whales in
the adjacent waters of Japan we marked the locality of the catch of eachindividual whale on a chart, wh:ch has led us to suppose that Sperm whalesin those waters l~ve in several different groups. So we have made our studyof their foetuses according to the four different sea areas, the Bonin
Islands, the Kinan, the Sanriku and the Hokkaido-Pacific sea areas. In
other respects, the method of our study is the same as in the case of Fin
and Sei whales; we have drew the growth curves of the foetuses of Sprem
whales by the average body-length per ten days and that per month.
In Fig. 4 you will find a growth curve of foetuses of Sperm whales in
the Kinan sea area though the cruve is imperfect for lack of material.
The curves in Fig. 5 shows the growth of the foetuses .in the Bonin Islands
sea area. The whaling season here lasts only three months,' ~arch, April
and May, and yet we can dearly discern that foetuses group themselves
along the two growth curves. Figs .. 6 and 7 show the growth curves ot
the Sanriku and the Hokkaido-Pac~fic sea areas respect:vely, either of whichshow two groups of foetuses with different tendencies.
Fig.

15

to

reet

4

_]___

126

Statistic Study of Foetuses of Whales

Fig.

5

Fig.

6

15

10

5

re et

127

Kazuhiro Mizue and Hisako· Jimbo
Fig.

7

-.

15

..

••

10

II>

..

5

feet

lb: . .

V.ar.

Apr.

Mey

Jun.

Jui.

Au._g.

Sep.

·, Oct.

Nov.

Dec.

month

Comparison of the curves of these four sea areas has proved that there
is no variation between the tendencies of each· curve. So, upon the combined data of these curves we have formed body-length distribution curves
per months. Then tracing the vallegs appearing in the monthly curves,
we calssified the foetuses into two groups, upper and lower. We have·
prepared a growth curve for each group of the foetuses based upon average
body-lengths per ten days and per month in the same way as before.
When we arrange the curve of the lower group to the left of the curveof the upper one we can complete the growth curve of the foetuses ofSperm whales. (Fig. 8)
According to this curve the pairing peribd of Sperm whales in ouradjacent waters is March. As for the body-length at delivery, Wheeler
made a report of a Sperm whale immediately after birth measuring 13.4 feet,captured in the neighbouring waters of Bermuda Island, the Atlantic, in .
1933. So did Bennett of the one measuring 14.2 feet in 1840, and reportedby Matsuura two measuring 17 feet. Figs 5, 6 and 7, foetuses measuring
13 feet are found in num?ers but those of 14 feet are comparatively few
while those of 15 feet are rare. From this we assume that the body-length

Statistic Study of Foetuses of Whales

'128

Fig.

8

•

!'eet.

l~~ ~en. ~r.
.. onthat delivery is 14-15 feet. The g~owth curve in Fig. 8 shows that the sizeof. 14-15 feet is attained to at August, so that the delivery time of Spermwhales is to be determined as August. However, this growth curve whose
composing members are rather homogeneous at first, grows in breadth andat the time of delivery it has the breadth of two months and a half. So
the delivery period of Sperm whales may practically be determined as
lasting from the end of July to the beginning of October.
If we are to determine the dura~ion of pregnancy of Sperm whales
from the delivery period shown on the curve of the foetuses, which isAugust, it lasts 17 ·months. The pairing period of Sperm whales differsfrom that of Baleen whales by two months, and the duration of pregnancy
here determined is much longer than theirs, which naturally brings about
a different delivery period.
(D) In this present study, we have determined the pairing period, theduration of pregnar..cy, the delivery period and the body-lengtl;i at del:veryof Fin whales, Sei whales and Sperm whales. Fig. 9 shows the distributionof the foetuses of Right, Blue and Hump-back whales in our material. The
growth curves coi:r:cic;le, with six months' variation; with those of Blue andKazuhiro Mizue and Hisako Jimbo

129

Hump-back whales in the Antarctic prepared by Mackintosh & Wheeler
and Matthews. The pairing period of Baleen whales in the adjacent watersof Japan and perhaps in the North semisphere at large is January.
Fig.

9Blu" <fheJ..,•

Hump t:ie.olt "l'he.le

Right l.tiaL

I

15

IQ

t"eet

l

.rsn.

•eb.

~--)

~ar.

TI

'llOnthIn Figs. 10, 11 and 12, you will find the seasonal distribution of foetus'body-length of Fin, Sei and Sperm whales in successive eight years.

Statistic Study of Foetuses of Whales

130

Fig.

I

1941

.I

I

1942

I

-~---

I

I•
/
/

>

· nonit_,,. /

~./
12
;

/

9

'

l?

I

1945

I

I

l.94f

I

I

----+.~

••

~

I

I
I

.•'

'·

I

12

'

1942

6

'

I

I

I

I

• I

./
I

I

J

9

;

12

/'

I

•;

I

I

/
6

9

12

'

ll

194,

I•

- - - - •7I - -

I
.;/

12
I
I

9

1948

I

I
I•

6.

-·

/

I

Fig.

1941

12

9

-~---;--

~

9

I••
/

I

/,

/

6

6

,1

I

~

;

I

I

t'eet

.;···
.

I

I

I

I•

I

tO - - - - ; - - -I

j_ ----r-I -

\947

I

,.

I

I

I

I

./

!'

,.

.,,.

.

I
/

I

I

I

1944

I

•j

~---;y

I

feet-

' I

..

I

----,--- -

-

I

194,

I

•/

10

10

; 1944

0

9

12

,.

·'

Kazuhiro Mizue and Hisako Jimbo
Fig.

131

12

/
/
10

-~-

.
···
.
•. .
:i~::

.;ci•"•. i:.
... ~··e .

_,,,,,,.

12'

6

l~

(Mar. 7, 1949)/

I'

Biological Survey of Fin and Blue \Vhales Taken in
the Antarctic Season 1947,...,48 by the Japanese Fleet
October 1948
byMasaharu Nishiwaki and Kazuo Hayashi
CHAPTER I. Introduction
I. Permission of Whaling for the Season 1947-48.
The whaling in the Antarctic water for the season 1947--48, a second
one since the cessation of the War, was granted by the GHQ, SCAP.
II. Objects o! the Whaling Survey.
The present investigations were carried out aboard the " Hashidatemaru '', a whaler of the Nippon Suisan K. K. (Japan Marine Products Co.,
Ltd.), and the "Nisshin-maru ", a whaler of the Taiyo Gyogyo K. K: (Taiyo
Fisheries Co., Ltd.) which went out on the permission of the above during
8 December 1947 to 10 March 1948. The movement of the vessels are•shown in Fig. 1. During this season, the whales caught were : 713 fin ·whales (Balaenoptera physalus), 608 blue whales (Balaenoptera musculus)
and 2 Sperm whales (Physeter catodon), of which 3 heads of fin were lostby the Hashidate Mam and were, therfore, not included in the investigation.
Sperm whales were also excluded from survey.
The chief object of the present survey was the classification of whalescaught in the Antarctic a,s part of the world-wide investigations on cetaceae.

On the other hand, this report was drawn up in order to present numericalappreciation of the Japanese whaling fleet. At the same time, data which
would serve as bases for the biological studies of the Cetaceae were collected as much as possible. In sliort, this is a statistical data and any
phenomena which might accompany them have been included in the
Appendix. · Physiology of various organs of the whales and anatomical
studies of the foetuses have been omitted.
III. Method of Work ·1) Body colour: Blue whales- Colour of the skin; size, clarity, number
and distribution of pale spots on the skin ; size, clarity and number of whiteflecks ; location of ventral grooves ; . clarity of striation ; were all observed
by naked eyes.

Masaharu Nishiwaki and Kazuo Hayashi-

133

Fin whales- Shading of the skin colour and extension of the pigmentation to the ventral surface from the back were observed by naked eyes.
2) Measuring·of the various parts of the body were made by steel tapeson those parts described by Mackintosh and Wheeler in their Discovery
Reports, i. e. :( 1 ) Total length.
( 2 ) Low.er jaw ; projection beyod tip of snout.
( 3) Tip of snout to blow-hole.
( 4 ) Tip of snout to angle of gape.
( 5) Tip of snout to center of eye.
( 7 ) Eye to ear (centres).
( 8 ) Notch of flukes to posterior emagination of dorsal fin.
( 9 ) Flukes, width at insertion.
(10) Notch of flukes to anus.
(11) Notch of flukes to umbilicus.
(12). Notch of flukes to end of ventral grooves.
(13) Anus to reproductive aperture (centres).
(14) Dorsal fin, ventral hight.
(15) Dorsal fin, length of base.
(16) Flrpper, tip to axilla.
(17) Flipper, tip to anterior end of lower border.
(18) Flipper, length along curves of lower border.
(19) Flipper, greatest width.
(20) Severed head, condyle to tip.
(21) Skull, greatest width.
(22) Skull length, condyle to tip of premaxilla.
(23) Flipper, tip to head of humerus.
(24) Tail, depth at dorsal fin.
(25) Flukes, notch to tip.
(26) Flukes, total spread.
Since (2), (4), (22) and (26) seemed to contain a large amount of measuring
errors, these were omitted in the present investigation, and following
measurements were added :(27) Breadth of the body, back to front (at tip of flipper).
(28) Height of the body when lying on its side.
(29) Greatest circumference.

134

Biological Survey of Fin and Blue Whales Taken(30) Tail flukes, from tip to tip.
(31) Lower jaw bones, length along curve of border.
3) The determination of the weight of various parts of whale body was
carried out with a platform scale, cutting the parts down to the size (~bout
50 cm3) to fit the scale and adding up the result. · Loss of blood and body
fluids had to be tolerated but small pieces sawed off were collected as muchas possible and also added up.
Investigations on testes1 overfes, mammary glands, food, thickness of
blubber, etc., were carried out by biologists apointed by the Japanese
Government and several men under their guidance so as to eliminate thedifference by individual fovestigation.
IV. Activities of the Japanese Whaling Fleet in the Whaling Ground(South of Lat, 60° S, east of Long, 90° E., and west of Long 170° W,)
granted for the Season 1947-48. ·Fig. i: The movement of Japanese fleets in Antarctic ocean (1947~1948 Expedition)

Masaharu Nishiwaki and Kazuo Hayashi

135

The Hashidate-maru Fleet left on 6 November 1947, and the Nisshinmaru Fleet left Yokosuka on the same date arr;ving in the ground on 5
and 7 December 1947, respectively. Their activities after those dates are
as shown in Fig. 1. The Nisshin-maru confined her activ;ties to an area
enclosed by Lat. 63° to 68° S., and Long. 155° to 172° E.,. but Hashidatemaru went further into the Ross Sea for operation. The present expedition
found that the opening to the Ross Sea was not in the direction of theBalleny Is. as usual but far to the east and the fleet found it difficult tofind the opening.
V. The Amount of Catch by the Japanese Fleet inside the Whaling Groundgranted for Season 1947-1948.
The Amount of whales caught by the expedition, is shown in Figs. 2
and 3 by month. As can be seen from these Figures, both the HashidateFig. 2. Catches of Blue and Fin Whales
maru and Nisshin-maru caught
Hashidate-maru fleet
many blue whales at the beginBlue wlval•- - -- Fin whalening of the season but theseanimals began to grow less by,,'
January and the fleets decided
tO go after fin whales. Nis'~shin-maru continue.cl to ope'''O/.
'
rate in the same area and1l~O·~=--~----r-~--o--~~---j.-----~~~-=-=-/-;
encountered the largest catch
DecemberJanuary
FebruaryMarchDatoof fin whales during February.
Fig. 3.
Hashidate-maru, however wentt~isain-Oaru rloet
into the Ross Sea and operated
100''
for blue whales. Fig. 4 shows
the amount of whales caught
according to different areas!' ' \
which is shown by the same'
scale as used in· the Whaling''
Ground of the Antarctic Ocean,',,v
6th Series (1) of Whale Resurce Data published by the"..
n•~OL--D-ee-~-bo_r_,_~-J-aru-e-ry~---.~-,,-h~bru-a-ry~~ ~rch Japan Association of Whaling
Dato
Industry.
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Biological Survey of Fin and Blue Whales Taken

136

Fig. 4. The map of catch on Japanese fleets (1947-1948 Expedition)+---+---+-----164

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VI. Material and Data.
The number of whales examined during the present investigations on
the Hashidate-maru and Nisshin-maru was as ·follows:Masaharu Nishiwaki and Kazuo Hayashi

r.reBlue Whale Female
Totel

Fin Whaler"

FemaleTotal

137

Hashidate·maru

Nisshin-maru

Total

138
136
274

200
236
436

338
372
710

87
124
211

176
221
397

263
345
608

Grand total of whales caught 1318 (1014 B. W. U.)

Investigation data on individual whales are to be published by the .
Marine Products Bureau aside from the investigation report.
Average time elapsed between the killing and treating of whales was
as follows:
Hashidate-maru

Fleet
--

Nisshin-maru·-----~~--------~~-

Species

Buie whale

Fin whale

Buie whale

Fin whale-------

Sex

Male

FemaleMale

FemaleMale

FemaleMale

FemaleAveragetimeinterval

71 '3611

71 061141 57 1151 471161 161151 51 1131 551141 2311

Unless otherwise stated, all data used in Figures and Tables refer tothe total figures and data obtained by both fleet, i. e. results obtained by
the Japanese Whaling Fleet.
VII. Abbreviations used.
Abbreviations used in this report and in the appendix are as follows :In the " Ossification of vertebrae " :1. Number and state of epi. in thoracic (or lumber) refer to the number
and state of epiphyses in thoracic series (or in lumber series) of vertebrae.
2. In the column for state, ank. refer to anklosed, those not statedrefer to " not ankylosed ".
On stomach contents :1. Size: L=Large, ea. 5.0 cm. and over (from rostrum to tail).
M =Medium, from ea. 4.0 cm. to 5.0 cm.
S=Small, up to ea. 4.0 cm.
X=Mixture of conspicuously different sizes.
? =Sizes indistinct due to high degree of digestion.
2. Quantity: O=Empty.
r=:;;Very small amount of krill
rr=Small amount of krillI

138

Biological Survey of Fin and Blue Whales Taken

rrr=Moderate amount of krill
R=Large amount of krill
3. Degree of freshness : f =Almost digested
ff= Half digested
fff=Fresh
F =Very freshOn the thickness of blubber :1. Point 1- The point on the horizontal cut side of the body (at the· position of lateral line in fish), where it intersects a verticalline from the dorsal fin.
2. Point 2- The point on the vertical cut near the earhole, where it
intersects a mid-dorsal line.
External parasites: irt=Much intectec;l or fully developed (diatOm)*=Moderately infected or partially devloped (diatom)+ =Scarcely infected or developed (diatom)
No notation=Not infected in naked eye.
White scar:
Number- O=None
l=Few
2=Scarce
3=Normal
4=Numerous
5= Very numerous
Colour of the blue whale : Pale spots, whits flecks and striation.
Number- O=None
State of distinction- !=Indistinct
l=Few
II= Distinct
2=Scarce
III= Very distinct
3=Normal
4=Numerous
5 =Very numerous
Colour of the fin whales :Body colour- N =Normal
B=Blackish
Extension of pigmentation on ventral groovesU =Upper(11-13 stripes of ventral grooves upward from navel line)

Masaharu Nishiwaki and Kazuo Hayashi

139

N=Normal
L=Lower
Tongue of pigmentation behind anus+ =Present-=Absent
Meeting of pigmentation in front of tail flukes+= Fused-=Not fused
CHAPTER II. Composition of Whales
I. Composition of Whales Taken.

The formation of whales taken by the present expedition is shown m
Fig. 5. The modes of blue whale male are 76, 78 and 82 feet while thatof female is 81. feet. Those in fin whale male and female are 67 and 71
feet, respectively.
Fig. 5. Size of whales taken (from all examples ofJapanese fleet in 1947-48 expedition)- - - - - Total curn1

60

II.

- - Melis curve- - - - - - Female curvo

S~x Ratio.

In blue whale, the sex ratio was 47 ..6% male while that in fin whalewas 43.3% male. The sex ratio of the foetuses was 48.8% male in blue
whales and 57.4% male in fin whales.

Biological Survey of Fin and Blue Whales Taken

140

Figs. 6 and 7 show the sex ratio according to body length. In blue
whales, there were about 60% of ma1e in whales of under 78 feet whereas there are none in those over 88 feet. In other words, male blue whales
do not grow to over 88 feet in length, at least, under the present inves- ·tigations.
In fin whale, the sex ratio of foetuses was 57.4% male (ea. 60%).
About 60% of whales under 69 feet were male, but the number dropped
suddenly in those over 70 feet in length.
The reason for this sudden fall in the number of males is that thefemales normally reach a greater length than the males, as is showngraphically in Fig .. 5, in which the number of whales examined is plottedagainst their length.
The whalers naturally select the largest whale to catch, which so often
happens to be a female so that the figures as a whole are not representativeof the whale population in that region. However, the sex ratio worked
out from data obtained on whales up to 78 feet in total length, which may
be expected to show the proportion of males and females more correctly,
show that the sex ratio is about 60% of males and not 43.3%, as would
appear if the total catch alone is considered.
These figures show the sex ratio among whales of similar length but· not necessarily of those of similar age. The proportion of males among
those of similar age is probably less than 90%. The males stop growing
after reaching a length of about 80 feet and consequently, are removed(

Fig. 6. Sex ratio and total length (Blue whale)

100

...~
'.;20

.

bt

io!,--~-=-~~~~~-~-;;,,--~~~-,,,,-~~;--~o,;--~-,,.--'~;;-o5

foot~•••

15

20

70

Total length in feet75

Whor.•

85

90

Masaharu Nishiwaki and Kazuo Hayashi

141

Fig. 7,. Sex ratio and total length (Fin whale)
100

~

c

~

!l
'O 20
1;<
m

!

~ 0 0L~~~~~10,.--~~15,._.........~20:--~~~5<5~~6oo-~65"'--'"-n,-_,_~
0;--~~~~

5

WhalesFoe tu sea

Total length in feetfrom the length classes of their contemporary females which grows to morethan 80 feet. The length classes containing those of 80 feet and less, must,
therefore, contain the males of an age equal to that of the females over
80 feet in length. Among whales of similar age,· therefore, the primary
sex ratio found in the foetuses is probaoly steadily declining from the highproportion of males to something nearer equality as the whales grow older.
III. Composition of Whales taken as Classified by the " International
whaling Statistics Board ".
The size of whales when reaching sexual maturity was taken from theworks of Mackintosh and · Wheeler, the blue and fin whale generally
reaching their maturity at the following length:Blue whale- Male
63 feet, 8 inches (63 feet)
65 feet, 7 inches (65 feet)
FemaleTable I. Composition of Blue whales
Classification

NumberRatio (%)

Group 1 (70 feet and less)Group 2 (71-85 feet)
Group 3 (86 feet and over)
Total

9
639
62
710

1.27
90.00
8.73

Immature male
Mature male

28
310

8.58
91.42

Immature female
Mature female

61
311

16.40
83.60'

Total No. of immature animal
Total No. of mature animal

89
621

12.54
87.46

·--···-

Biological Survey of Fin and Blue Whales Taken142

Fin whale- Male
Femal_e
Table II.
Classification

74 feet, 2 inches (74 feet)
77 feet, 9 inches (77 feet)
Composition of Fin whales
NumberRatio(%)

Grot,lp 1 (55 feet and less)Group 2 (56-65 feet)
Group 3 (66 feet and over)
Total

0
110
49S
608

0
'18.0981.91

Immature male
Mature male

6
257

2.28
97.72

Immature female
Mature female

13
332

3.77
96.23

Total No. of immature animal
Total No. of mature animal

19
589

3.12,
96.88

GHAPTER III. Composition of Male Whales
I. Sexual Maturity of Male Whales.
The sexual maturity of male whales are determined by the following :1. Size of the penis
2. Size of the testis
3. Weight of the testis
4. Histology of the testis.
The present investigations took the weight of the testis as a chief
indication of maturity with additional considerations on other points.
II. Relationship beween the size and weight of testis.
In their Discovery Report, Mackintosh and Weeler classified male
whales according to the size of the testis ("the size of the testis represented'by a number obtained by multiplying together the length, breadth anddepth measured in centimetres").
Ih order to obtain the· relationship beween the size and weight of thetestis of male whales taken by the Nisshin-maru, various parts were plottedagainst body length and the average values obtained are .shown graphicallyin Figs. Sa, Sb, 9a and 9b. Figs. 10 and 11 show the comparison of theirsize and weight. Naturally, these Figures show that both weight and size
curves have the sa~e tendencies and seem to show that the weight method
in which values taken from fresh ones c_an be used, is much more exact
than the size method which necessitates calculatiqns for tabulation.

Masaharu Nishiwaki and Kazuo Hayashi

143

Fig. Sa Size of testis in different length ,(Blue whale) on Nisshin-maru No. 1

88

84 .
80

76
72
68

64
60

Fig. Sb Size of testis in different length(Fin whale) on Nisshin-maru No. 1

56

52
45

~..

.
...

44

90
88

8

84

a
76
28
72
24
68
64

56
52

A\·.

48

44

r

OL-6=9~7=0~7~1~72~7=~~7=4,--;75~~76~~77,..--7~8~7~9_,,8~0.....,,,8~1-8~2---,8~},--;8~4-,;8~5-:::86~8""7
Length of wh!lle in feet

40

56

r

52

l

28
24
20

16

~12
.. 8

\

~

(!; ..
r.
. . .."

r'

w

.

'ci
~ 4

~0

>

•.

62

6~

65

06

Langth of whole in reet

67

6e

69

70

71

72

n

74

144

Biological Survey of Fin and Blue Whales Taken
Fig. 9a Weight of testis in different length(Blue whale) on Nisshin-maru No. 1
&l

76

72
68

64
60
56
I

52

..
i

48

.
0

.,
0

44

40

;.

}6

"

}2

28

~

!1

2

h

a.

~20
76

.

:i

68

...

"'

0

7l 72 7j

7

0

i

60

~ 068 69 70

Length of whale in feet

52
4
4

·}2

..
I

I

... .r .. '.
". •r ,;r
r i
I

I

'I

.;,

lo

..

\

:0

:"

r,

"' 16
!i

.

.

~l

0

~ 8

Sc

0

.:; 4

.!1
:Ii 0

~

~ 12

64

24

f

; 16

72

28

"'

0

~

61 62 6} 64
5t.engt.h of wha~e in feet.

8

69

70

71

72

Fig. 9b Weight of testis in different length(Fin whale) on Nisshin-maru No. 1

7}

74

77

78

79 80 8L 82

8} 84

85

87

Masaharu Nishiwaki and Kazuo Hayashi

145

Fig. 10 Size of testis in different length of whales
andon Nisshin-maru No. 1
Weight of testis in different length of whales
6

Volume curve-· Weight. curv1t- - - • Volume curve frOtD Discovery

R.e~orta

5

r-~/

4

/\

/

I
I

I
,,,_-.... ,..__--,

/

n n n

n ,

\\

\

,__..J~
9 ~1l.el\S~l> of wllale ln foe~ (Blue whale)

\

n •

-,,

I

w ~ m ~

'

\

'\
'\

\

',

'I

~

M

I

\

~

Fig. 11 Size of testis in different length of whales·andon Nisshin-maru No. 1
Weight of testis in different length of whales
6

,,,,,

.

.J- .3.
....
~--- - _,

.. -

--

~

1

0

i

~

:

Voluaie ouJ"Y'e
We:la,h'\ '\.\Jrve

0

!

~

0 :

~1,...__,,2,.._....,.~:--.......,....-----,~5~-,,6~"""T...--"Fa,.......--,6~9~~1Mo~-:;;n~-:;:n:----=n:----=r.---=~~
Lengt,h ot vhal~ in fee~ (Fin whale)

III. Weight of the Testis.
The 'weight of the test;s of all whales taken by the Japanese fleet in
the present whaling season were taken to observe overall tendencies.
As shown in Fig. 12, the development curve of testis in blue whales.
is very gradual at first but becomes rapid when the body length reaches.
73-75 feet, and again becomes slower thereafter. In male blue whales.

146

Biological Survey 'of Fin and Blue Whales Takenof over 83 feet in length, decrease of sexual· function can generally beseen.. This point constitutes one of the b:ggest difference from the curveof the number of ·corpora lutea in female whales. These tendencies
observed in the present investigations coincldes well with the reports of
Makintosh and Wheeler as found in p. 406 of their Discovery Report, Vol.
· 1. This evidence is shown by the average curve taken from their Fig. 133(p. 406) plotted on· Fig. 10.
The same tendencies can also be seen fin whales. As can be seenfrom Fig. 13, the develoment curve makes a sudden upturn around a bodylength of 62 feet and takes . a natural course of development thereafter.
The fact that a remarkable decrease· phenomenon cannot Qe seen must be
due to the failure to catch large, aged whales.
Fig. 12 Weight of ·testis in different length (Blue whale)
BO

72

64

60

•

.. -.. - -

52

I

48

9

~

,

-•

t

9

II

..
.
•.. ..

..:

I rr

28

::

1'
~

20

'

'"'

; 16

..

¥

12

:

8

0

.. 4

!a
::

0

I

•
I

8

0
66 6 7 68 • 971 72 7} 7
Len~t.h of whale in feet_

I

•

75

7')

0

77 7i 79 80 81 82 BS 84 85 86 87

'. J'

Masaharu Nishiwaki and Kazuo Hayashi

147

Fig. 13 Weight of' testis in different length (Fin whale)
76

72
68
64

60
56

5
48
44
40

,6

,2
28

f

...

!.
•

~

24

......

~ 16~12...i 6

..

0

~ 4

!;1

~

5

6

67

68

9

70

71

72

7'

IV. Sexual and Physical Maturity according to Body Length.
The maturity of males according to their body length is shown in Fig.
14. The sexual maturity was classlfied only by the weight of testis as isdone and not by macro- or microscopic observations of the testis. If the
combined weight of both testis was less than 10 kg in blue whale and 5 kg
jn fin whale, they were considered immature.
The physical maturity of tnales was considered complete when all theirepiphyses of vertebral column were fully ankylosed. They were considered
immature if thoracic epiphyses were not ankylosed even if caudal andlumber ones were.
Considered from these points, the sexual maturity of Blue whales is-reached when the body length· is about 74 feet, and the physical maturity
at about 80 feet: The sexual maturity in Fin whales is reached at about
63 feet of body length and the phys'cal maturity at about 70 feet.

Biological Survey Of Fin and Blue Whales Taken

148

hrcsntage of' maturity

..

"'

0

8.

t>l

,

Perc&ntage of maturity
t>l
fj

°'

0
°'

0

0

fj

. :;

~,..

~

•
•

.

~

'""'

V. Average Length of Whales. according to the Weight of Testis.
Fig. 15 and 16 show the average length of whales according to theweight of testis from which it can be seen that in blue whales the bodylength of whales having 10 kg testis is about 73 feet but the length
suddenly incr~ases to 76 feet in those having 12 kg testis. In fin whales,Fig•. 15 Average length of whales in each class of weight of testicles (Blue whale)

eo

2

6

10

14

18

22

Weight of testicles in kg.

26

~

. :)4

,6

42

"6

'50

'54

,8

62

70

74

78

82

Masaharu Nishiwaki and Kazuo Hayashi

149

Fig. 16 Average .length of whales in each classof weight of testicles (Fin whale)--------

the' curve for whales having testis weighing up to 6 kg take a sudden
upward trend which then becomes gradual. The body length of whales
having 4 kg testis is about 63 feet, and that of 6 kg testis about 66 feet.
The foregoing is a good evidence that the classification of maturity bybody length is based on the classification by testicular weight.
· Figs. 17 and 18 show· the curves for average· testicular weight at
dJferent body length according to physical maturity. The physical maturity
curve in Fig. 14 thow the ratio of the mature and immature in the former
data. These figures also furnish good evidence of the various factors as
stated above. It seems correct to observe that the body length of male
whales at its maturity, as seen from Fig. 14, is 79 feet in fin whales.
Fig. 17 Average weight of testicles in different lengthon physical maturity (Blue whale)

.,

,1
I

"°

1'~
I

/,
/

""-,

I

/
\I

...

..
0

~

;; 1

•

-,

',

I

I

I

I

I

I

I•

I

'

'

' ,__
Me't.ure

150

Biological Survey of Fin and· Blue Whales Taken
Fig. 18 Average weight of testicles in different lengthon physical maturity (Fin whale)//______-... _______ ,,. ,,,,."'.... .........

Mature/--~-------

-

/
-/

m
~

u

~
~

m

..
3
0

~ 10

:§>
~

5'
:;

~ OL,.,,.~~~~~~~~~~..,...,-----,~___,.,~-=-~-=-~,,,---;;;--,;:~
~
Q
M
~
~
~
~
~
~
9
~

n

n

n

Body length ln feet

CHAPTER IV. Composition of Female Whales
I. Factors decisive in Determining Sexual Maturity of Female Whales.
In order to investigate the sexual maturity in cow whales, following
points must be considered :1. Presence of a foetus
2. Presence of c?rpora lutea in the ovaries
3. Size of the uterus
4. Size of the ovaries
5. Weight of the ovaries
6. Condition of the ovades follicle
7. Condition oi the mammary glands.
However, the present researches took chief account in the presence ofcopora lutea in the ovaries with addit:onal considerations on 1, 5, 6 and 7.
II. Relationship between Body Length and the Number of Corpora Lutea.
Figs. 19 and 20 show the distribution of the number of corpora lutea
according to body length. In Blue whales, the curve is gradual up to bodylength of 79 feet (unc~r 3 corpora) but takes a sudden upwar:d swing after
it reaches 80 feet in length. The same is seen in Fin whales, the curve
being gradual .up to 65 feet body length (under 2 corpora) but becomessuddenly high at 66 to 67 feet length.
ill. Average Length of Whales according to the Number of Corpora lutes.
Figs. 21 and 22 show the curves for average ler gth of whales accor-

Masaharu Nishiwaki and Kazuo Hayashi

151

ding to the number of corpura lutera observed. In blue whales, the average
length of body for those having 3 corpora lutea is 79 feet but those having
4 corpora suddenly goes up to 82 feet. In fin whales, those having 3 corpora
lutea is 68 feet and those with 4 corpora, 70 feet. From Figs. 21 arid 22
it can be seen that the average body length according to the number ~f
con:ora lutea is 84 feet in blue whales and 72 feet in fin whales.
Inother words, the standard body length in female blue whales is 84 ·feet.and that in fin whales, 72 feet.
IV. Relationship between Sexual Maturity and body Length.
F. g. 23 shows the percentage curve of sexually mature (possessing
corpora lutea) animals according to body length from wh:ch it can be seenthat sexual maturity in blue whales is reached at about 78 feet and thatin fin whale females at 67 f~et.
I

Fig. 19 Length of whale and numberof corpor lutea (Blue whale)
41

29

27

2}

21

19

17

l}

11
e

!l

~

::

9

7

f'

~'

.c#li. ••Q88lb&.

08:.oal

Ji

••

s, 0

co 0

~•a•

j h-~-=~:'!!:'.!:~•-a_•~•-•_+~~~~~~~~~-::--='

67

9 . 71

7}

Lej,gt)l ot vhale in f'tllei

75

77

79

81

8}

8'}

87

89

91

9}

9'

97

Biological Survey of Fin and Blue Whales Taken

152

Fig. 20· Length of whale and number.
of corpora lutea (Fin whale)
''4'

'iii

,7

"
''
~l

29
27

-.

2,
21
19
17

•9

!

.:l

i
g

.-........ .

'I!

.:

~

.50

,2

,4

56

58

~t.11 of whale. in feo\

60

62

Fig. 21 Average length of whales in each class of corpora lutea numbers (Blue whale)
88

87
86

~81

·;so
.

~ 79

=.1e

5

:11'-~~~~-5~..,..~.,,.--.,.ll,.....~l~,..,...~15,.....~1~7~~1~,......,,,,~~2,,....._25.,.__2=7,.....~2~9~,~l~~,~,~,=5~~,~7~,=9,.....~4-1~4~,~
Number of corpora lutee

153

Masaharu Nishiwaki and Kazuo Hayashi
Fig. 22 Average ltjngth of whales in each class of corpora lutea numbers (Fin whale)

72

71

I
'

9

11

1,

15

19

17

21

2,

25

27

29

}l

"

Number of corpora lutes

Fig. 23 Maturity s;urve on different ' length (Female)
100

60

Phy Bica l ma tur 1 t7

Body length 1n feet (Blue whale)

Sexual maturit'6o

60
Body· length in teet. (Fin whale)

65

70

'

75 ;

,5

'7

,9

I

,

Biological Survey of Fin and Blue Whales Taken

154

Fig. 24 shows the classification of female whales, indicated in thecomposition of whales taken, according to sexually immature, pregnant,resting and lactating by their body lengths.
In blue whales, the height of curves for the immature and the mature(resting and pregnant) are at different places. The trend of the curves forresting and pregnant match well and show that· approximately half of themature whales are pregnant.
In fin whales, there are a very small number of immature femalesdue, probably, to the fact that whalers generally aim to catch large sizes.
Approximately one-third of the mature females are pregnant and the trendof the curve by their body lengths is similar to the above.
Fig. 24 Maturity of female in all Japanese fleet- - - Immature--·------- PregMnt- - - - R.esting
Lactating:
0

20

~
~

"'g

zo1...-..::.,,,~,..-~~.L.....:~,,-~~~~o~~~--.eo..,..-.,--~~~~.--~~~~~~~

l.ength of whale in f'eet (Blua whale)
ImmatureI

ReU.ing'0Lactating

I

20

~

__ /

.!!

~10

.I "\ .\ .

.

Pregnant.

\

,_./
I

'

\

·--,
,.---··, __ -. ,'\\\

Ii

_,"

_,

\\
\~, ,."''

0,,..._ ............

~~•
io'---~"""'~~~~"'-~"-'""""".__~~---'-:=-·~---•---·~-=-~~~~-=0

57

60

10

o

eo

Length of whale in f'eet (Fin wh11le)V. Monthly Ratio of Pregnant Whales.
Fig. 25 shows the monthly ratio of pregnant whales and endorses theviews of previous workers in the l?regnancy rate decreases monthly from
December to March. Following figures show the pregnancy rate as a whole:
Table ill
Whale speces

Pregnant

Resting

LactatingBlue whales

162
53.29%

Fin whales

32.17%

131
46.38%
210
66.88%

1
0.33%
3
0.95%

101

Total Mature
Female304
314

Masaharu Nishiwaki and Kazuo Hayashi

155

Fig. 25 Percentage of pregnant whales in the catch by month(contain the whale, which has function corpora lutea & no foetus)-

Tot.aJ.

- - - Nhein-ma-ru- - Haahideta--maru

.

i.

.

~

e
!

,,..

~5

/

~

.......l.
'~0£0'-~Da~c-em~be-r~.1------.Ja-~-•=ry~~-----.F=eb=ru=ar~y~-+-M~a~rc~h~
Data

100

s

.

!::.

f10
~

2

...:;.='
l0ioa--:--....~t-----.--~~-+--..,-...,....,.
__~~..-,,-.,......
0.o•ber
February
March
Jenua.r~

Dow

VI. Relationship between the Number of Corpora Lutea and the Largest
Graafian Follicles.
Figs. 26 to 31 show the results of observations on the relationshipbetween the largest Graafian follicles and the number of corpora lutea.
Due to the time allowed for handling, this work was carried out on 236
hea~s of blue whales and 221 heads of fin whales taken by Nisshin-maru.
Figs. 26 and 29 show the diameter of the largest Graafian follicle ineach animal_ and the average in each body lergth class. Figs. 27 and 30
give the relation. of the number of corpora lutea according to the bodylength (same as Figs. 19 and 20 but only on data from those taken by the
Nisshin-maru). Althogh they fail to show, at first glance, any presence ofa relationship between them, their average curve alone plotted on the same
diagram show, as in Figs. 28 and 31, that there is a tendency of coinci-

,4,
i

41
2001

,9

I

,7,

,5

''

~
E-<

,1

150

rn

Q)

c;;
.r::

::::
Q)

...

::l

P'.l
"Cl

100

§

19

.si.«

ri

17

'S
»

~

~

~

. ...

e
~

~ 501

'

..
•

~
~

:

..

.. ..
-

-

ii v..

.

g30.

70----nLength ot whe 1e t n teet.

78

~

&l

~ &i

82

""
~

0

o

. . . . ......
;

a

84

c

.

<;;I

~

8 a :

~G

.....

0

:

0

:g

11

t:

.. . i-/.·. ~·

-.:
!

Q

l'

: ~

g

~

I

15

~ }i
:

0

Q

~

G

11
M

:E

88

90~2-

9

Fig. 26 Diameter of largest Graafian follicle(Blue whale) on Nisshin-maru No. 1.

~6i- 6971

7,

9i- 9,

9} - 97

Length of whale in teetFig. 27 Length of whale and number of corpora lutea(Blue whale) on Nisshin-maru No. I.

Masaharu Nishiwaki and Kazuo Hayashi

157

Fig. 28 Average diameter of largest Graafian follicle and
Average number of corpora lutea on Nisshin-matu No. 1. (Blue whale)
Diameter aurvs
Number curve

60

o

20

i

//

1--/
I
/
/

:10
:
~ ~

:

~

/1

I

I
\
I

I

I

/

\/

__ , , ,. """' ',II

o
~8

I

I

69

70 71

72 7,

74

78

79 80

81

2 8} 84 85

86 87 88 89 90 91 92 9}

Length or whale in'" feetFig. 29 Length of whales and Number of corpora Iutea (Fin whale on Nisshin-maru No. l.

41

}1

29

27

25
2}

21

19

17
15
1}

..

11

81 •

.
~

0

~ 5

...
0

~

il,,-,,..,.._..,.~~~~~,..-:,,,...--,,~~:-=---:::;.---;;::--.::--;;;:ao
50

52

54

56

5a

(.etlgth of whale in feet

o

10

12

74

76

78

aa

158

Biological Survey of Fjn and Blue Whales Taken
Fig. 30 Diameter of largest Graafian follicle (Fin whale) on Nisshin-maru No. 1.

l5

50

.:
a, IB

...
....

II. ...

0

! 0
Q

•••

"-~56,-~5~8~760""""~6~2~674,...--~6~6~6~8,,.-~70~~72~~7~4~~76..-~~~80..:....~
Length of whale in teetFig. 31 Average diameter of largest Graafian follicle and
Average number of corpora lutea on Nisshin-marn:_No. 1. (Fin whale)60

Diem&ter- curve;'\

Nurcber cur~•

''
' ''
I
t
•

.

50 ;o

I

,.

u

~40

·'

.
c

I

...

I

I

I

...
0

...

l :
II ;
:;;" 10 2.... ..

,>-,,-/,//j! <0~~~~~~6r69~~~-=-1
56 57 58 59 60
Length of whale in feet

61

62

6~

~4

65

65

67

6•0

69

70

71

72

75

74

75

76

77

78

Masaharu Nishiwaki and Kazuo Hayashi

159

dence. In other words, the appearance of the largest diameter in Graafianfollicles appear with the increase in the number of· corpora lutea, i. e. withadvancing age.
VII. Frequency Curve for the Number of Corpora Lutea.
Figs. 32 and 33 show the frequency curve for the number of corpora
lutea the peaks for wihch in blue whales occur at 2, 6, 9, 12, 15 and 19(dotted lines show the value g:ven in the Discovery Report). These peaks
in fin whales occur at 2,, 5 7, 10, 14, 18 and 21, which coincide well withthe peaks for fin whales of 1, 5, 7, 10, 13 and 15 obtained by sumrnariFig. 32 Frequency of numbers of corpora lutea (Blue whale)

60

Frequency cune of thie expedi'Uo1o---- Freouency curve of Diacovery Aepor'Le

..
0

I\
I

\
\

10

·I

!

g

\

'·

,
'
0
~ -·~o~~...,.-5..-~~,.10.--~-,...-~-~---"~,_::.c:.--"''--~~=--""='?,._--=::::;:~;........,,,c.~i~5Mtacb.er of co-r901'e lut.ea

Fig. 33 Frequency of numbers o(corpora lutea (Fin whale)- - Frequency curve ot t.hte expedition- - -- Frequency curve of Diecovery Report•

c

g,20

·g
~
c

~

g10

~·

f
~

I

I
\

,-~

v'

~ 0 '-oo--~;~~-,,10:;----~15-=----:~_:_L-....~~__\L--:::~~_...,;~~....L.:::;:,....
Nuitbr.r C>f corpora lutea

160

Biological Survey of Fjn and Blue Whales Taken

zation of various values obtai:ped for Antarctic fin whales. These curves
for fin and bltie whales show that, especially in fin whales, there aresmall number of comparatively aged whales with over 13 corpora lutea.
On the whale, there are more number of aged animals which seerp.s toindicate the fact that the whaler go after larger animals when the composition of whales in general is getting younger.
VIII. Relationship between Sexual Maturity and the Weight of Ovaries.
Fig. 34 Average weight of ovaries on different length of whales (Blue whale).

10Pregne-·

6

,,,

\

5

/

/ ....

4----____,,.

.... ,

II'/ a..~1'ng
\

I

I

\t

,,.,,,,/

I
I

I

,
•
!2
. !ii

,

/

I
' ../ I!.
r·--_..,.,, . . . . . ,_,,,..../·:1-·-·-·-·-·-·-·--.
.
.
_,
·
...

__ .... ,

/"--

.,

. . . . . . . Sexual immature, ,!f:11 oi..,,,,,4-=5,....,,,,.....,.6,,..7..,..,.a.,,,6""9.,,,70=-=-71-=12_,,,7},.,..,,,74:--:75,,..-,1"'"6..,1""1-.1=a--1=9_,,,oo,--:ca1"""""'a2::--::a=-,"""a""4'""'e;=-e.s=-"'a1,...,,,aa....,,,a9~90=""""9""°1""'9"'"2'""'9=J.
Wngth of whale ln feet.

Fig. 35 .Average weight of ovaries on different le~gth of whales (Fin whale}

.•
'
I

I

I

I

,,
,,,.---:.. ___ / /Pregnant --~~~e_t~r;_g__,/'-·-·-·-·-·-·-·-·

-·-·

-·

---·'
,/...........-4;"""'·;/\./.........../

/

''

I

\ _ _....--·- Se:rual immature....... ,'Masaharu Nishiwaki and Kazuo Hayashi

161

Figs. 34 and 35 indicate the weight of ovaries in sexually immature,
restirig and pregnant whales classified according to their body lengths.
This coincides will data funrnished in the Discovery Report and needs nofurther comment.
IX. Average Number of corpora lutea according. to Body length.
The number of corpora lutea averaged by body length in physicallymature whales is shown in Figs. 36 and 37 (Cf. Figs. 19 and 20), Fig. 25
shows the phys~cal maturity curve given by the percentages of the above.
From these figures, it can be said that the body length of physicallymature animals is about 85 feet for blue whales and about 71 feet for fin
whales.
Fig. 36 Average number lutea indifferent
length on physical maturity,~oMature

,,~
1

/\

I

/· .........1

I

,'

\

..........

\..,./

I

I

:

~ I

t

l I

'" ~
I

\/

(
Immature

Length of whale in feet (Blue whale)

Fig. 37 Average number lutea indifferent
length on physical maturity

I Mature
I
,\

I

'<
\

.[

/

,,...............

IF--.,,-v,/".,' '

'•"

I

'./

I

'
•

111111.e. ture

~ oL~:::,,,.,,.~~~::::::....~_:_~,.--~~~--:==-~__:~--;;;;~
6?

70

I/

~

Length of whale in feet (Fin whale)

X. Mammary Glands.
The present investigations were carried out on the colour and thicknessof mammary glands on 372 heads of blue whales and 345 heads of Fin
whales, values taken being the average of them all. Those class"fied asBiological Survey of Fin and Blue Whales Taken

162

lactating are animals from which even a drop of milk was observed during
treatment, other~ being classified as "not lactating." The latter was again
divlded into pregnant, resting and immature according to the condition of
the ovaries; e. g. those possessing functional corpora lutea classed as the" pregnant " (functional corpora lutea of pregnancy and of ovulation were
Fig. 38 a Average thickness of mammary glands in different length (Blue whale)
'14

17

I}

16

12

15

11

14

o hetating

10

,,
/

''

v

/

''

'

''

Prf'li<;nant.

.
.;1

~ 3

~'!i 2 '~o'...
~-ng;<;6;'.7i~oii"f'!han1-1.1l12•1n773•r.'.1,7:47"75:--7.76r-i7;.;7-77lia-,,79'><aoin""".a;;>1-0:a2:--08:;-5-,8;;;:4-;;;e),--.,1:6;,--,8"°'7'""""'88;--;8"'°9....,90=-""91~9-2_9_}_94-

Fig. 38:b '. Average thickness of mammary glands in different length (Fin whale)
21J
l}

20

28

12

19

27

11

JS

26

10

17

9

16

25
Lacte. ting

14

211
2}

15
72

n

22

____ ,.,,---- .-Lerigth of whsle· in feet

Masaharu Nishiwaki and Kazuo Hayashi

163

not classified), those possessing no· corpora lutea in the ovaries as the" immature " and others as " resting ". Figs. 38 and 39 show these
averages according to body length. It is noted that both the immatureand other curves increase with the increase in body length while that for
lactating is in a place by itself with no relation to the former.
There are always a great variations in colour of mammary glands but
they have been averaged as follows :As a whole, the colour of mammary glands in young whales is paleI
which increases in tone as the whales grow older. In aged animals, thecolour becomes deep brown. The colour oi mammary glands in lactating
whales is orange tinted, irrespective of their age. This tendency is in
keeping with the fact that the thickness of mammary glands and that ofblubber always change irrespective of the age of whales.
Following table shows the percentage rate in colour of mammary glandsin blue whales.
Table IV. Colour of mammary glands in blue whales ·Nisshin-maru

I
Immature

No.

%

10
28.6

RestingNo.

2%Pregnant

No.

2.1
2
2.1

Lactating

No.

%

Pink

White

Ivory

7
20.0
3
3.1
2
2.1

7
20.0
3
3.1
5
5.2

I. Cmnamon.
I Reddish
Yellow [ Tawny
i

I

I
:

6
17.1
22
22.9
23
23.7

i

3
8.6

r

!

25
26.1
22
2,2.7

I

!

%

i

-1

I

I
I
I
I
II
I

i

2
5.7
34
35.4
38
39.2
1
100

I

Brown

II

I
I

!
I

7
7.3
5
5.2

i

Hashidate·marui---"---~-----

I

Immature

No.

Resting

No.

l

%

5.9
2
3.0

%
I

Pregnant
Lactating

i

No.

%
No.
I

I

%

I

i

Pink

Cinnamon

Tawny

Brown

15
45.5

I

3
9.1
4
23.3

I
II

10

15.2

29
42.9

1
3.1
9
52.9
11
16.7

3
17.3
14
21.2

White

i

14
42.4

I
I

I

Following table shows the percentage reate m colour of mammary
gland in fin whales.

164

Masaharu Nishiwaki and Kazuo Hayashi
Table V. Colour of mammary glands in Fin whales

Nisshin-maru
White

Pink

Ivory

13
62.0
5
3.8
1
1.6

4
19.0
4
3.0
2
. 3.1

3
2.3

----

immature
Resting
Pregnant
Lactating

No.
%
No.
%
No.

%

1
1.6

No.
%

Cinnamon I Reddish
I Yellow Tawny
I
4
I
19.0 I
46
40
3134.6 i
30.2
23.3
I
26
17
15
I
42.6
23.5
26.6
I
1
100.0
i

I

Brown

4

3.0
2
3.1
i

Hashidate-maru

Immature

No.

Resting

No.

%%

Pregnant

No.

Lactating

No.

%
%

While

Ivor~_J Cinnamon I

4
44.5
2
2.9
2
5.3

1
11.0
9
12.9
6
15.8

I

I
I

I

4
44.5
35
50.0
14
36.8

I

Tawny

Brown

14
20.0
10
26.3
2
100.0

10
14.2
6
15.8

CHAPTER V. Foetuses
Main growth curves for 162 heads of blue whale foetus and 101 headsof fin whale foetus, classified according to the date their mothers werecaught and by body length, were taken. There were nothing new to belearned from them, only endorsing the data contained in the Discovery
Report. The monthly me.an body length on the curves, especially, coincidedwith the older data.
The sex ratio of the foetuses is as described in previous chapter. On
the whole, there were 48.8% male in blue whales and 57.4% male in fin
The smallest foetus was thatwhales and 57 .4% male in fin whales.
measuring 10.6 inches in the blue whale and 7.9 inches in the fin whale,
while the largest obtained was 22' 211 in the blue whales and 15' 3" in fin
whales.

165

Biological Survey of Fin and Blue Whales Taken
Fig. 39a Mean growth curve of foetuses(Blue whale)

Fig. 39b Mean growth curve of foetuses(Fin whale)

18

22

18

17

21

17

15

20

15

19

15

·.

15

/.forch

e
14

14

.

I}

..

1:;

12

12

ll

11

...

10

9

10

•.

••

.. .. . . .
.....
:

8

7

•

0

.. ..., .

9

®

..
.. ..

GQ

•

\

•o

...

O•o • o•.,

•.

i }

e

~

o,

.

0

o

GD

•o

••

..

0

~

•

.
0

.,o

i! 2

.r.

·.

5

••

~ It:•~

0

0 .....

., 5 •

8'

8

~

6:

. ..

""

1

o inHvidusl foet1,.19ea 't• ·inttividuel. foetuaen ~• eversgs lenzth in month- mean growth curve'O•.:; 1 01l'

~ o~~~~~~~~~~~~~~
Deeeimber
Da~eJanuary

February~Qrch~ O'--~~~~~~~.,-,--~~~~~J e nusry~'ebrusry
Me.rchDecember
Date

CHAPTER VI. External Characters
I. Colour.

There are many reports on the colour of whales. Observations on thecolour of blue whales during the present expeditions followed the notations
made by Mackintosh and Wheeler on pale spots, i. e. " most of the bodyis covered with a pale mottling which cons:sts of small, roughly oval marksof colour which is similar to, but lighter than the blue-grey background",on their clarity and number, and on white flecks on which the two authors
say that " those pale spots may also be seen here and there on the ventral. grooves ". The clarity on the appearance of striation on the ventral surfaceof tail flukes was also shown by percentages. These results are shown in
Table VI. · The name of the fleet was noted · in because of the fear that

Masaharu Nishiwaki and Kazuo Hayashi

166

personal view may differ in individual cases.
Table

VI.

Pale sport
Sex

FEMALE

MALE

Fleet

Hashidate-maru- --Gradingof Nos.

No.

_,~

-

i

;

%

No.

!
)

0

0

i

-

0

1
2

I

3
4

7
17
58
44

39
2
40
84

5

9

52.0
12.6
43.0
32.6
6.6

0
40

I

I
I

Hashidate-maru

Nisshin-maru

%

No.

No.

!

19,5

0
4

0
40

I

1.0
20.0
42.0

52
43

8
45

ii 19.2
I

17.5

6

93
48

Nisshin-maru

-~-

35

I
I
I

!

i
:
'

'

26

I

%

!

3.1
19.8
39,7
32.8
4.6

%

18.1
3.4

I
I

i
I

39.8
20.5

Clarity
0

0

-

I
II
III

19
34

16.1
28.8
55.1

15

I

-

139
21

20.0
69.5

0
10
24

10.5

87

I

I
I
!

-

0

8.3
19.8

29
177
28

71.9

I 12.475.6
12.0

White Flecks
0
1
2

3
4
5

I

I

I

0
17
39
48

12.7

27

20.1
2.3

3

29.1
35.8

i
I
I

I,,
I

I

0.5
35,5

1
71
13
35
55
25

4

'8.5

0
20
26

15.1
19,7

17.5
27.5
12.5

53
24
·9

18.2
. 6.8

1.5

0
42

40.2

1.7
26.8
6.4

63
15
64
48

21.2
20.4

41

17.5

Striation
0

I
II

0
36'
43

III

55

26.9
32.1
41.0

i

I

!
1

I

I

3
64

I

I I

102

• 32.0i' 51.0

39

I

31

I

53

I

l

15.5

I

31.3
29.1
36.6

4

1.7

75

31.0
48.9

115
41

17.5
'

On the colour of fin whales, Mackintosh and Wheeler says in their
Discovery Report, that " the most obvious f eatu re is that pigment covers
the whole of the back and flanks, while the ventral surface remains
unpigmented ". This shading of body colour over the back and flanks(not darkened due to light and air), extention of pigment over the ventral
surface, as follows, were examined :i) Extention of the blackened area of the back over the ventral grooveas though by brush;,ii) Extention of the blackened area of the back and flanks in tonguelike form towards anus from the tail ; andiii) Whether the blaekened area of back and flanks meet form either

167

Biological Survey of Fin and Blue Whales Takenside just in front of the tail flukes.
Table VII shows the number of whales in percentage on above points.
Table VII
MALE

Sex

FEMALE--------·-------------~-------------------!
Hashidate-maru
Fleet
Nisshin-maru I Hashidate-maru------ - -----i
Classificationi No. ofa-No:~ - - o- . i ~N~o-.-o-f--%-00____________ ! whales%
whales%! whalesNormal colour 'Blackened colour ·

67
15

81.7
18.3

154
22

87.5

!

83

12.5

i

38

68.6
31.4

Nisshin-maru-No. of_ _ _ __whales

%

205

92.7
7,3

16

49
27.8
25
20.2
High
13
16.3
15-1
36
58.1
58.0
50
102
61
49.2
58.S130
Norm.
of Pigm.
26.8
25
14.2
38
30.6
24.9
23
55
Low
T_o_n_g_ue-of-1~+-C-- _7_0__ ,_____81-.4--'---1-02_ __,__92-.0 ---+---8-9-----'~-72-.9_.L.___1_83-- ---8-2-.S--

Extention

Pigmentation
beh. anus
Meeting of \Pigmentation +in front of
flukes ·
J

16

18.6

14

8.0

33

27.1

38

17.2

73

85.9

99

56.3

90

73.8

110

49.8

12

14.1

77

43.7

32

26.2

111

50.2

As a result of foregoing data, the normal ·body colour of whales canbe summari;;;ed as follows :In blue whales, pale spots do not appear collectively but is dispersed;
there are 6 or 7 spots, normally, of a size about (4 x 6) cm 2 in an area of 1 m2 • The clarity of these spots is distinct. The white flecks are also
dispersed in an area about 1/3 behind the ventral grooves. The striation
is normally distinct and there can be seen no difference between sexes.
In fin . whales, the normal body colour is slate-gray, -and the extentionof pigmentation to the ventral groove is on the lltli or 13th from the navel
line, neither higher nm lower. The tongue of pigmentation behind anus
is normally present. · The pigmentation also normally meets in front of thetail flukes. Here, also, it is hard to distinguish the. sexes according tobody colour·.
Although the distinct white flecks seen in blue whales seem to be due·to old age, the difference in body colour due to age is very slight.
II. Proportion of Body Length.
At the time of the weighing of whales on the Hashidate-maru, measurement of body proportion was also carried out (Cf. Table IX). According
to Mackintosh and· Wheeler, the proportion ol anterior part of the body
becomes larger with the increse of body length, both in blue and fin
whales. However, since the measurements made were small and the range

168

Masaharu Nishiwaki and Kazuo Hayashi

narrow in the present investigations, no calculations were made on themean values and their distribution according to body length group. There
were evidence, however, of the proportional. increase of anterior portion of
the body and decrease in the posterfor portion with the increase in age,
not necessary with body length. From what measurements that weremade during this expedition, no distinctive relationship can be seen between
the body length and its proportions.
From the relationship between the number of corpora lutea and body
proportion in female whales, as shown in Fig. 40, it can be seen in Nos.
3, .5 and 6, especially in 3 and 5, that the proportionate percentages of .
anterior portion increase with the incre~se in the number of corpora lutea.
In the measurement of posterior portion, as in Nos. 8, 10, 11 and 12, theproportionate percentage apparently decrease with the increase in thenumber of corpora lutea. (Numbers used here refer to the investigation
Nos. as explained in the "Method of work" in the Introduction).
III. Proportions of Body Weights.
Total weight of foetus was meas 1red 0:1 30 heads of blue whales and
16 heads of fin whales. Foetuses were cut into blocks of about 30 to 50.. kg. each and weighted on a platform scale of 200 kg. capacity. Some of
the body fluid has naturally been lost dudng the process so that· the weightcannot be taken as the true weight.
The relationship between body length, L (ft.), and body weight, W(kg.), obtained by the least squares, from all the measured, values on blue~nd fin whales, is as shown in Fig. 40a, the regression line in thesesamples being
1

Lz.ss

· W= 3.083 .......................................................... ( 1)for the foetusess of blue and fin whales.
Weight of variously dissected parts were we'ghed in a box placed on
the said platform scale on 4 heads of male and 10 heads of female blue
whales and 2 heads of male and 3 heads of female fin whales on the
Nisshin-maru, and on 7 heads of male and 9 heads of female blue whales,
and 4 heads of male and 7 ·heads of female fin whales on the Hashidatemaru (Cf. Table X). At the same time, oil production of these various
parts were measured by the readings on oil tank gauge from part of thesematerial. Since the total weight of various parts could not be obtainedBiological Survey of Fin and Blue Whales Taken

169

Fig. 40a Weight-length relation of foetuses (Blue & Fin whale)

due to loss of body
fluids, excreta, urine. and stomach contents,
overall weight of a600
whale could not be
500
taken. If on ·the other
hand, the proportion}00
of were the same inwhales, their relationship to each other0
would be similar to100
the total body weight.
800Fig. 40 b shows the
7c60
total weight plotted
50
accor.ding to bodylength.
}0Although the process of dissection of
the two fleets diffeted
slightly, there would"'~10
be no effect on the~~~~4~~·~5~-o-~~-8.,.--9~10~~~~~~~·
Lo; L (kg.)
measured values withthe exception of one or two polnts. In grand total, especially, same items
have been inculuded on values from both fleet.
Total weight : Measurements were made on 30 heads of blue and16 heads of fin whales. Their regression line, measured as for foetus,
obtained from the specimens by the least squares, are
~J~

L3.5

Blue Whales ...... w= . x104 ........................... ( 2)
5 27

.
L~.9
.
Fm whales ......... w = .SOxl02 . . . . . . . . . . . . . . . . . . . . . . . . . . . ( 3)4

, CHAPTER VII. Thickness of Blubber· I. Sites of Measurement ·Thickness .of blubber was measured in following places:
Point 1-- The point on the horizontal . cut side of the body (at the170

Masaharu Nishiwaki and Kazuo Hayashi
Fig. 40b Weight·length relation of whales (Blue & Fin whale)

13
140
1;0

60

66
Loa L (foot)

68

70

z2

74

76

78

so

e2

84

86

88

90

lO(l

position of . lateral line in fish), where it intersects a verticalline from the dorsal fin.
Point 2- ·The point on the vertical cut near the earhole,- where it
intersects a mid-dorsal line.
II. Average Thickness of Blubber according to Sexes and Body Length
measured on the Nisshin-maru Fleet.
Figs. 41 to 44 show mean values measured by the Nisshin-maru Fleeton the thickness of blubber according to sexes and body length. Figs. 41
and 42 show curves for male whales and indicate that the thickness
approximately increases in proportion to body length. It is interestingthat in· blue whales, thickness becomes less with age when the animals
attain sexual maturity. Curves in Figs. 43 and 44 represent those forfemales which peculiar modes appear every 3 feet in blue whales andevery 4 feet in fin whales.
III. Monthly Change in the Thickness of Blubber.
Average percentage monthly change of the th;ckness of blubber bybody length are shown in Figs. 45 and 46. From these Figures, it can beseen that the thickness of the blubber increases during December to March,Biological Survey of Fin and Blue Whales Taken ·171

Fig. 41 Variations of thickness of blubber in length of whales (Male Blue whale)
14

l~

12

I
I

I

----'

'

I

°O 10

.
G

·~

'

',

,/

',

"',

,,,----,"i'i_____ ·~.\- - - - - - - -/- - .... ~"'- - - Point 2
,
,,..
/
~ 9
''./'/ ',,JI
G,',,r-----.J-'-.:::::.,~ -- -· -

- - -

- - -

- - - - - mean linos""

! a.,.,,_..,,,..-=--=,,._=--=--=.,--=--::--=,...._....,....~~~~~~~~~---.~~~~
9 N n n n
~ n
- n M ~ oo m m ~ M m ~ Q M.l..ength of Whalo in feetFig. 42 Variations of thickness of blubber in length of whales (Male Fin whale)
10

Len.g:tt. of whale in feet·i. e. the summer months in the Antarctic, irrespective of the sexual stateof the whales. In other words, the state of nutrition of the whales
become better during these months. Th'.s is particularly remarkable inblue whales.
Due to the lack of whales in lactating stage, no curve for such animals
was obtained but this has been noted in places. As can be seen from the
Figures, the thickness is remarcably small which seems to have a relative
connection with the fact that the thickness of mammary glands in lactating
whales is conspicuously thick.

Masaharu Nishiwaki and Kazuo Hayashi

172

Fig. 43 Variations of thickness of blubber in length of whales (Female Blue whale)

12

c

~ 11

v

~ 10

/\

f\.

- - - -·-/

/

...-

/'.....
'\

I/. . . . . . . . . .
"-

I

',

~,- - - - - - - - - - - ~- - --/~- - ~~/·----- \,._;__ - - --~, - - - - mMn lina
\

,'

//\

\

Point 2I\/ \\ ... --'

/
\

',,

/
\

I

~J

/

\

',

/
\

J

\'

'/
Leng;th of .whale irl foet . .

Fig. 44 Variations of thickness of blubber in length of whales (Female Fin whale)10

V j overage

I

'\; 7

.... ....----\

/\

,' \
I

/,,,.
\

/

______ -· __ - "\"" - _ ~ - - - - L - - - ~v~ i

Point l

/

\

/

/

60

61

/

2

6~

4

5

66

7

-

-

-

I.

V

- - - -

- - !1tei-ag1J

76

78

//

...,

I
\

\ ___ ..1

'

-

,'

I

\

/

\

58 ~ 59\/\

Point 2 ,,."'_,

I

/

\

/

,- ......

// \

', /

I

"

68

69

10

n

12

n

74

n

77

,

IV. Relc:itionship between the Thickness of the Blubber and Body Length
as measured on the Hashidate-maru Fleet.
The coeficient of the thickness of blubber is shown by D/L x 10 where
D denotes the thickness of blubber in cm., and L, the body length in feet.
Distribution of its mean value at Point 1 (at the side of the body) is shownin Figs. 47 and 48. In general, both the male and female blue whales
coefficient of from 0.09 to 2.0, the peaks being presentat 3 points between
1.1-1.5. In Fin whales, the coefficient is distributed between O. 7 to 1.5,
peaks being present in 1 to 3 places between o.9 and 1.3.
V. Relationship between the Thickness of Blubber and Body Length by

173

Biological Survey of Fin and Blue Whales Taken
Fig. 45 Monthly average thickness of- blubber (Blue whale)
Point 2

16
Point. ll?
14________ .,...

.cl

Saxu:ll irnma ture

.....

1;

~

.

~ 12

12

~ ll

11

~ J.0

lO

•

-------~etureSexual immature: 9"-~--~r-------i----~-~Doc ember
Januaryf'ebrunry
March

DecemberJanuary

February

March

Pregnant

16

I
/

I
..

l6

I ,
14

-- -Deoember,.......,.. • .......-'-_,,,.,.-::_·_

Ja rlUary

--

14

,
/ Sexual,/.

1

Pregnant.

/Resting 15

i;il'llmoture

1

12<sexual//inm:a ture

11

-//

February

Januaryo.aF{;brua.ry

Merer

o. 7

Fig. 46 Monthly average thickness of blubber (Fin ~hale)
Point 2Poini l

12

12

il

..,

__ -

10

9

DecemberJanuary

FebruaryMarch'"_ ... - - Doc ember

January

il'ebruary

Merch

DecemberJanuary

Februery

March

15
14

12
11

11

10

10

0.6

0.6

o. 7

Doc ember

January

February

March

Date

the Degree of Sexual Maturity, as measured on the Hashidate-maru.
The di~tribution of mean v~lues as shown in Figs. 47 and 48 can bedivided into several groups by the mature and immature 1n males, andinmmature, pregnant and resting in females. These are as shown in Figs.

Masaharu Nishiwaki and Kazuo Hayashi

174

Fig. 47 Tickness of Blubber (Blue whale)......_____.

Maloo- ---oFem.:.le

R
'1
I

•

I

I

I

I
I

10

I
I

"' "- -·
'

I

/
I

I

!.._

I

'

I

\l

Fig. 48 Tickness of Blubber (Fin whale)
Mele
F'ema.J•o-----o
20

10

~

'\
b._
'O..

}

0 0

90 .

100

110

120

1}0

' 'O..

140"--- ?"S.-o-

150

160

-0

l 70

180

49 and 50 which indicate that in immature whales both male and female,the average value is distributed between 1.0-:-1.4 and seems to have auniform mode, although the trend is not remarkable probably due to the.
small number of whales measured. The mode in mature males is at 1.3,
especially that in whales from the Ross Sea being at 1.5, showing clearly
the difference in the thickness of blubber. The distribution is broader
both in the pregnant and resting, although the distribution in the pregnant
In both cases, whales from the Rose Sea
whales is slightly higher.
contains larger proportion of higher coefficients.
The coefficient for immature fin whale males is situated around 0.91.0. There is no remarkable difference in females according to month, but175

Biological Survey of Fin and Blue Whales Taken
Fig. 49 Tickness of Blubber (Blue whale)
Re.at..ing

Immature.. ~--o.
i·Dea.Jan.
it.oee-eea(Feb.)

10

10

5

•0

.

'.l

00

100

140

120

100

160

200

0,

M

200

20

0

.a

i•

Pregnant
Dec.Jan.

<>---<> Rose-l!lea(Feb.)
· 10

10

Fig. 50 Tickness of Blubber (Fin whale)l11111ature

Resting~--oo----·-oMeloFemale10lt---l<

Dec.
Jan.
Feb.

10

5

140

'l·ec.

Mature male

o---o

Jali.

lf---« Feb.
10

0

60

lOC

m the pregnant and resting whales larger coeficients increase as themonth advances. On the whole the difference in the coefficients between
blue and fin whales seem to be essential.
CHAPTER VIII. Food
Krills and Other Fish
The present investigation also disclosed the fact that the chief food of
the blue and fin whales in the Antarctic Ocean is the krill. Of the 710

I.

176

Masaharu Nishiwaki and Kazuo Hayashi

heads of blue and 608 heads of fin whales examined, fish other than krill
found in their stomach were as follows :Fish of large Harpodontidae: 2 cases (1 in blue whale, 1 in fin whale)
Squids: 4 cases (1 in blue whale, 3 in fin whale)
Small type of Harpodontidae: 24 cases (10 in blue whale, 14 in fin
whales)Kind of Iniomi: 1 case in blue whale.
Jelly fish : 3 cases in fin whale.
II. Euphausia as Whale Food
Results of investigations on Euphausia, especially E. superba, are shown
Tables VIII and IX, in which the empty spaces are mostly those in which
the krill was found almost digested as reddish-brown colour in the intestines.
It is interesting to note that the semi-monthly percentage rate of stomachs
with krill coincide wholly with the catch curves as shown in the foregoing
Figs. 2 and 3. In other words, th:s shows how the whales move theirpositions according to the presence of krill or that the investigation of krill
would enable judgement on condition of the catch.
Table VIII. Stomach contente (Blue whale)
"Cl

:-a
..:.:

Q)

('!)

.....
0

Half-months

::i::
Ill
::r

December~JanuaryUl;5.:..,iIll3'Feburary

i;::

::ii

('!)
('!)

March

......

~
Ul::r

December

Ul

s·
..,~
i;::

January
Feburary

::ii

('!)
('!)

......

March

]

,.c:

-5;:l

:c..:.:

"CJ

~

,.c:

.c:

Q)

I

,.c:

I ,.c:u

u

ro

s

tl

......

I

·c

I

x

0

~

('!)

s I ·~,.c:

+->

Q)

8

g

·§ro I ,.c:

>,
P.

I

z

Ill

·~

,.c:
u

ro

ro

i

u

s I s

s

0

0
+->

~

I :a..:.:

tl

ro

I

0
+->

......

b

~

b
:::::

~

~

s

I:;'

b

I .:t:..,...,.c:

·c
..:.:

I

'

I

·c

~

!

..:.:
'<-<

I

0

....>,

I

Q)

P.
>,

~

f-<

\

'<-<

0

'El

~

·~

d

z

c5

z

I
z I :?R

.c:
·~

d

z

d

z

d

z

zd

~)~jui)x)~!

41
79
27
39
36
52

26
53
12
11
23
31

15
26
15
28
13
21

. 63
66
44
28
64
60

7
3
0
0
1
3

3
15
1
3
10
9

2
17
4
3

14

-

-

-

-

-

112
78
66
87
31
38
24

99
58
83
51
16
13

13
20
33
36
15
25
15

88
74
50
59
51
34
37

44
27
3
10
1
3
1

-

Ul

9

-

'El

Ul

'<-<

0

0

-

-

·~

-

.~

I

!

10

I 2

7

12

5 14
21
- 8
- 8
2 10
8 14

-

-

-

-

-

24
10
9
11
7
6
4

14
11
9
17
2
2
3

17

j -

I --

18

I 57

10

12
13
6
2
1

--

-

--

-

1 5
3 26
- 4
1 2

-

24 41 31
5 19 33
1 8 24
5 11 35
1 6 9- 1 17- 4I 5
-

-1-

1.
3.

- i
2'
1

9

8

-

3 1 -· - -

'

_I_

-

- .
'

-

.,

Biological Survey of Fin and Blue Whales Taken

177

Table IX. Stomach contente (Fin whale)~~g,Half-months

If.....
'Sj'z0
DecemberJanuary
Feburary
March

'S0

z0

z

5
47
44
43
53
1

2
22
19
23
25
0

3
25
25
20
28

13

4

141

-

40
47

I

43
54
47
0
22
-

1-~~~~~~~~-i-~----C---

DecemberJanuary
Feburary

18

16

2

67
44
56

38

29
16
28

143
63

28
28
85
35
5

58
28

89
57
64
50
59
56

1

5

7

4

3
4
1

6
5

4

6

2

10
9

1

-

1

8

3

7

2

1
1
4

8
9

1
2

7
10

2
l

8

1

11

1

2

-

---~-~--~-"-1-~~-1-~- 1~ ~
6
8

-

2

2

-1-

1~

2
9
10
10
14

2 -

i =

6
9

6
2

2620-4915-,---

14

=

20

I 25

26

9
3

I 1_·11

1_0
1

6

22

56

1

_

~ I 14 21 _ _
1
83
6
March
- 1- _: ..:1!1=
I--'------~--'---~---'---~-~-~---'---~--'--'---'---'-----'
Symbols used in tables are as follows :L=E. superba. Large, 5.0 cm. and over (from rostrum to tail).
M=E. superba. Medium, from ea. 4.0-5.0 cm.
S=E. superba. Small, up to ea. 4.0 cm.
X=E. superba. Mixture of conspicuously different sizes.
?=E. superba. ·Sizes unknown due to high degree of d:gestion.
R=Stomach with large amount of krill.
rrr=Stomach with moderate amount of krill.
rr =Stomach with small amount of krill.
r=Stomach with very small amount of. krill.
CHAPTER IX. Prasites
I. External Parasites
Following external parasites have been collected from blue and fin
whales apart from certain more or less minutes form found on the baleen :Cirripediae- Coronula sp. (C. regina)·Conchoderma sp.

Masaharu Nishiwaki and Kazuo Hayashi

178•. · Coppepodae- Pennella sp.
Anphipodae- Cyamus sp.
Diatomes- Cocconeis sp. ·The incidence of these parasites are shown in . Tables X. and XI.
Semmrtally, there is no great changE; according to whaling seaosn with
Cirripediae, but Pennela is found more in the earlier part of the whaling
season, one example being a clue whale on which about 30 full grownspecimen had been found hanging. Cyamus was found in the latter part
· of the whaling season. Almost all of the diatomes found were of the
Cocconeis sp., which began to increase, both in incidence and density, as
the whaling season advanced.
Table X. Percentage Incidence of External Parasites in Blue. Whales.
Infected
NisshinHashidate
Maru
Maru

I

Whale lice
(Cyamus Sp.)
Barnacles(Coronula Sp.)
Conchoderma Sp.
Parasitic coppepedae(Pennella Sp.)
Diatomes

17
7

~

'i

1
2
72

'

I

I

32

Not Infected
I Hashidate
NisshinI
I Maru I Maru
!
176
402
!

Percentage of Infection
NisshinHashidate
Maru
Maru
i

6.32%

7.37%
3.92%

430

2.60%
I 0.37%

426

0.74%

1.84%

325

26.76%

25.12%

i

17

417

262

4

268

8

267

109

l97

i

I
I

I

I

0.92%

Cpndition unknown: 2 whales used as fenders in the Nisshin-maru Fleet.
5 whales missed from the Hashidate-maru Fleet.
Table XL Perceqtage Incidence of External Parasites in Fin Whales.
Infected
NisshinHashidate
Maru
Maru
Whale lice
(Cyamus Sp.)
Barnacles(Coronula Sp.)
Conchoderma Sp.

24

19

24

30

1

4

Parasitic coppepedae(Pennella Sp.)
Diatomes

1

4

77

131

Not Infected
Hashidate
Maru

I

184

!

I

184

:

207

:
!

I

207

I

131

!

Percentage of Infection
NisshinMaru

Nisshin : -Hashidate
Maru
Maru
378

I

11.53%

4.79%

367

I 11.53%

7.56%

393

I

0.48%

1.01%

I

\

iI

393

0.48%

266

37.02%

I

I

I, .

1.01%
3~.00%-

Conditions unknown : 3 whales missed from the Hashidate-maru Fleet.

The incidence of white scars was as · shown in Table XII whichindicates that no whale is free from it.
The proportion of scars obtained in the prevlous year and prior to it
were found to be, in the maiority of cases, m()re numerous for the older. one, the values being :179

Biological Survey .of Fin ana Blue Whales Taken

About 92% had more scars· obtained earlier than the previous year ;about 2% had equal number of scars obtained in the previous year andearlier, about 6% had more scar:s obtained during the previous year.
There were no difference between male and female, or between blueand fin whales.
It was thought possible to tell the age of whales by the white scars
but there are some remarkable exceptions that they would not constitutea very good facter in assuming age. There is a tendency, however, oflarger number of scars in aged animals.
Table XIL White scars on Blue Whales.

Name of fleet

Hashidate·maru

Classification

Numberof whale,
0 I
I
5 '
40 '
25 ~
57
6

none
few
scarcenormalnumerousvery numerous

I

Male

Sex

I

I

Nisshin·maru
Number [of whale.

%
o.o
3.8
30.0
18.8
42.9
4.5

0
39
2
40
84
35

I

%
0.0

I 19.5
II

1.0
20.0
42.0
17.5

FemaleNisshin-maru

Hashidate-maru
Numberof whale
0
6
46
27
51
4

I

I

i

i

Number Iof whale

%

0
70
8
26
90
41

0.0
4.5
34.3
20.2
38.0
3.0

I

I

i
I

I

'
i

I

%
0.0
29.8
3.4 .
11.1
38.3
17.4

Table Xlll. White scars on Fin Wha1es.
Male

Sex

Female--·-------1---------------1-------------Name of fleet

Hashidate-maru

Nisshin-maru

Hashidate-maru__c_1a_ss_ifi_c_a_tio_n__ 1-~-t-~_~_;_fel __%_0__ 1_-_:_u_~_~_;f_e.,I __%_o__ ,_~_fu_~-~--~-rel ___-·-~onefew
scarcenormalnumerousvery numerous

O
0
36
13
32
4

iI

,

0.0

O

0.0

0.0
42.4
15.3
37.6
4.7

58
5
14
81
18

33.0
2.8
8.0
46.0
10.2

I
I

:

25

I
!

26
30
45
15

l

Nisshin·maru%_ ~N~u-m~b_e_r~l--%--0.0
4.1
21.5
24.8
37.2
12.4

of w603hale:,i--o-.o-

I

9
27
108
28

I
··
!

I

26.8
3.9
11.5
45.9
11.9

II. Internal Parasites
Internal parasites are found more commonly than the external Crustaceae and they are often present in great number in whales. However,since not all of the intestines were cut open, no percentage of incidence
were obtained. The species found were tapeworms (Tetrabothrius sp. ?) or
Acanthocephala.
Numerous encysted tissues, due . probably to nematode worms, were
fuond near the body cavity, especially in the ventral muscles. Since calcu-

180· Masaharu Nishiwaki and Kazuo Hayashi

lation of its incidence would be difficult, no attempt · was made to obtain
percentage rates.
CHAPTER X. Summary
The results· of various observations made the catch by the Japanese
fleet during the season 1948-1949 can be summarized as follows :1) Composition of Whales Taken
There is no denying that the body length of whales taken is decreasing year by year. However, as far as the present expedition is concerned,
the results showed a slight tendency to improve in this respect. This isan evidence that the Japanese whalers have come to recognize the fact
and are taking great care to catch large-sized whales. It is regrettablethat four of the blue whales taken were under-sized and is therefore inviolation of the international law. However, it is hoped that everyone willrecognize the fact that the Japanese whalers are doing their best to comply
with the international code of ethics. Average body length were as follows:
Male
FemaleAnimalBlue whale77 .~6 ft.
81.09 ft.
79.55 ft.
66.52 ft.
70.53 ft.
68.80 ft.
Fin whale2) Body length of mature whales ·according to their sexes came out asfollows:
At sexual maturity :MaleFemaleBlue whale74ft.
78 ft.
63 ft.
68 ft.
Fin whaleMale
FemaleAt physical maturity :Blue whale79 ft.
85 ft.
68 ft.
Fin whale74 ft.
3) Pregnancy rate, especially in fin whales, during the present expedition
seemed to be lower than the precwar records, values being:
Blue whale- 53.29 % of total mature animals.
Fin whale32.17..% .of total mature animals.
4) It has been found that it is impossible to tell the sex of a whalefrom its body colour and that only a very inaccurate data can be obtained in telling the age of whales from their body colour.
5) Althe!'ugh it is still uncertain, judging of age from the condition ofwhite scars and the amount of parasites on whales would be a good

Biological Survey of Fin and Blue Whales Taken

181

subject of study in knowing migratory habit of whales.
6) It is to be regretted that no investigatiol1's were made on thebaleens.
APPENDIX I.
Number of Whales in the Ross Sea
I. Blue Whales in the Ross
. Sea
During the present season, the Hashidatemaru operated around theRoss Sea (Lat. 74°-76°S, Long._ 172°--178°W) from 11 to 25 February 1948.
During this time, 77 heads of blue whales alone (Nos. 393 to 471) weretaken and distribution of their body length is shown in Fig. Al. As aresult of investigations, there were no immature whale among 47 headsof males and only ·one immature whale among the 30 heads of females.
19 of the mature females, about 66%, were pregnant, 10 or 34%, resting.
Thispercentage
Fig. Al Length frequency curve on Hashidate·maru
Mele-

Wh!tles tekeri

f'rom Dec./47 t<> Jen./48- - - - Wht1le!J· taken from Rose-eeD

10

.

,,'

...

:2 5

I

I

'

\!

0

\

\
I

BoG..y length in feet

Feroa le

10

.

"
0

'

L

D

a
j.

' ' ,______ _'V<-Q---7~6~77~78~7-9~80~8-1_8_2~8~~8-4-~~-M~8-7_8_8~89~~~9-l
Body length in feetseems to be rather
high for ·February.
The ratio of maleto female was 100
males to 64 females,
incidence of males
being remarkably
higher than in otherwaters.
Averagebody lengthofwhales taken in
the Ross Sea was
78.66 ft. for males,
82.77 ft. for females, the total average being 80.62 ft.
Th's is slightly
above the average
but the percentage
of aged males wasMasaharu Nishiwaki and Kazuo Hayashi

182

Fig. A2 Length of whales and Number of corpora. lutea (Blue whale)41oWhales taken from Dec./47 to Jan../48•

Whales taken frolf! Rotrn-eee

,9

,1
29
27

25'

21

19

17
15
l}

11

.

9

.

~

.:: 7

~- 5

~

& •

.,

~ 1
i:Oo·

:ii ·'·

0

a. 0

.. -

... 0

0

8c.&dll~a ...... 1;i

11.

5

67

69

~ngt.1'1 ot~ whale

7l

ln feet.

7;

75

77

0

•o;i

79. 81

8}

85

87

~9

91

9}

higher than in other water, individuals in which ossification of vertebral
column had been ankylosed was 55% in males and 20% in females.
Immature animals were few in both male and female. There were alsoa very low incidence of external parasites, Cyamus being found only in
one case each in male and female. There was no evidence of Coronulasp., one each in male and female of Pennela sp. Diatome film infection
was found in 15% of the male and 30% of the female. As was explainedin the Chapter on the blubber, its thickness per unit body length waslarger the whales being fatter.
This was not only seen in external
appearance but also borne out by the fact that the production of 0:1 per
B. W. U. during operation in the Ross Sea was extremely high.
No particular difference from other waters was seen in the relationshipBiological Survey of Fin and Blue Whales Taken

183

between body length and number of corpora lutea (Cf. Fig. A2). This isan evidence that there were a small number of large-sized whales in theRoss Sea during the present operational season when considering the facts
that the average body length of whales taken in the same region by the
2nd Tonan-maru in 1941 were 79 ft. for males and 82 ft. for females, inaverage, and that a school of quite a large-sized blue whales had beentaken in the Kerguolen sector during the season 1923 -:- 1933 (A. H.
Laurie: Discovery Reports. Vol. XV, pp. 223-284). However, due to alack of biological data, no explanation can at present be given.
APPENDIX II.
Copulation of Humpback whales
There has been many tables told by whalers on the copulation ofwhales but very few description has been given by biologists. As far as
is known by the auther, the only report of such detail is the one by D.
C. Lilie published in 1910 (Cf. except at the end of this chapter).
During the present expedition, the auther witnessed following actions
by the humpback whales.
On 13 January 1948, at about 21-00, ·operation was being carried out
on the Nisshin-maru drifting about 1 mile off the so-called pack line.
There were many cracks and inlets into the pack ice near the ship andmany humpback whales (about 15 to 16 heads) were swimming aroundthere.
As though these animals know that the catching of humpback whalesin the Antarctic Ocean had been prohibited by the International agreement,
they often came very near the ships. Just then, some humpbacks started
to come nearer to the Nisshin-maru, making tremendous spashings. At· first, one stood watching without any thought but there were something
peculiar in their movement and the author began to watch with intent.
There were two heads ·of humpbacks and, as shown in Fig. A3, theyat first swam in single file. Shortly later, as shown in Fig. 4, the two
whales began to roll and romp together with each one of their fippers
well above the water. At one time, as shown in Fig. A5, one of thewhales flipped its peculiar tail flukes above the water and then dived deep
into the water, followed immediately by its mate. For a few seconds afterwards, the surface of the water remained calm and then the quiet of the184

Masaharu Nishiwaki and Kazuo Hayashi

water was broken by the sudden surfacing of the two whales vertically,as shown in Fig. 6, with their ventral surfaces in close contact and. appearing above water almost to the middle of their bodies, just below their flippers. Then the whales fell back into the water, sometimes together,sometimes. separately. After that, the same movements started all overagain. Although there were some pauses, the romping continued well overthree hours from the time it was first noticed until the whales were lost tosight. It may be that the objective had not been attained, but the so-called
Fig. A3 Swimming together

Fig. A4 Female turns one side, then male turns
another side and makes several dashes

Fig. A5 Dive into the depth

Fig. A6 Refloat vertically they faced their ventral sideBidlogical Survey of Fin and· Blue Whales Taken

185"standing up on the flippers", as shown in Fig. A6, were witnessed morethan to or three times. The author was of the opinion that this was the
copulation of humpback whales, the fact of which was borne out by theexperienced crews of the ship. The author had heard previously from awhaler that the whales jumped above the water, their ventral surfaces
together, during copulation but was very doubtful if this were feasible.
However, the present expedition showed that this is possible although the
copulation itself may be performed underwater. The whole scene was a
grand s'ght if slightly garish. It was a great pity that the time was at
sundown and did not permit photographing. The· nearest that whales cameto the ship was about 200-300 metres. Some whalers said that the surfaceof the water became was not witnessed at this time. It seems that thewhales moved away before the acts were completed but there were noevidence that the male whale heaved a great sight or become exhausted
after the act. Even during copulation, the whales rested a while, sometimes swiniming quiently for 15 to 30 minutes.

ABSTRACT
Observations on 'the Anatomy and· General Biology of Some
Members of the Larger Cetaceae
by D. G. Lilie, Hutchinson Research Student of
St. John's College, Cambridge.
(Taken form the Proceedings of the Zoological Society of London, 1910)

VIII.

Miscellaneous Observations

3. Copulation, period of gestation and rate of breeding.
The Balaenoptera are said by whalers to copulate at the surface of thesea. The pair swim towards each other and turn slightly on the sides sothat their ventral surfaces face one another. The male makes several
dashes at the female to insert the penis. · When the pair first rush together,
the long axes of their bodies are paralel with the surface of the sea ; but
they curve up vertically at the end of the act. After copulation, the maleis said to be exhausted and easily caugt.

Masaharu. Nishiwaki and Kazuo Hayashi

186

APPENDIX III.
· Age of Whales and Colour of Crystalline Lens\

Wheeler's statement in his Discovery Report says that " the determination of age on whales is a- question of much scientific research andeconomic importance ".
During the expedition, I h9-ve stood at the side of the slipway watching
the eyes of whales as they came up. Some looked as if they werepeacefully sleeping, whilst others had a hunted look. Amongst these, thecolour was of a golden shine. But I understand the colours vary.
When I was a boy, I used to be to told that there were witches (anold cat, an old fox or an old badger) who could change themselves !nto
men, then some hero would have to go out and dispose of them. These
Fig. A7 Comparison of body length, Weight of testicles
and colour of crystalline lens (Blue whale)overte~ro.e•vertebrae not enkyloe-?d(p!1yeical immature)o"lr.l"t"!~:."'ee helf(to lum·<)\""r t<<Jriee) ankylo9ed

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Biological Servey of Fin and Blue Whales Taken

187

animals (wiches) were always old, I was informed, and their eyes shone abright golden colour.
Looking at the eyes of whales, I wondered if this story could not be
connected with them.
In dissecting, first, cut the vertex corneae in a V shape mannerwithout scratching the lens and then take the lens out with the corpus
vitreum with two fingers. Secondly, wash the corpus vitreum and the iris
off with fresh water. It is then ready for observation in the shade. It is. best to look at the lens in. the shade and not in direct sunshine (though
the sun in the Antarctic was not strong).
Lastly, I have found at the crystalline lens, some to be yellow,· others
light yellow, and still others without colour or transparent.
It was interesting to discover that the colour of the eyes had a very
dose resemblance to that shown in the chart. There was no colour by
Fig. AS Comparison of b'ody length, Weight of testes
and colour of crystalline lens (Fin whale)
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Masaharu Nishiwaki and Kazuo Hayashi

188.

naked, eye, in the lens of the eyes especially, if the epjphyses of the lumbar series was not ankylosed. As ankylosis progressed, the colour became
yellowish, and when the ovaries had a number of corpora lutea (much as30) and complete ankylosis had set in, t~ey were found to be completely
yellow. · Dep~nding upon individual whale, the gradual change of colour
was also noticed with the number of corpora lutea or weight of testicles
and state of an.kylosis.
In order to gauge the change in colour, it would have been an advantage to have used the lovibond tintometer (the British Drughouses pattern)or some electric method, but I came upon this instrument too late to obtainfull advantage of its use.
The classified colour of lens in the chart has beeri done only throughThe five colours as classified are no colour (transmy naked eyes.
parent), coloured, light yellow, yellow, total yellow.
Fig. A9 Comparison of body length, Number of corpor lutea
and colour of crystalline lens (Blue whale)

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189

Biological Servey of Fin and Blue Whales Taken
Fig. AlO Comparison of body length, Number of corpor lutea
and colour of crystalline lens (Fin whale)

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The data gathered in this Report may be of some help in determining
the age of whales through the colour of the crystalline lens.
It is hoped to include colourimeter and electric ammeter m the next
expedition in order to complete this research but the present report will
only suffice by preliminary descriptions. (Cf. Figs. A7-A10).

•

Masaharu Nishiwaki anc1 :Kazuo Hayashi

References
N. A. Mackintosh, A. R. C. S. and J. F. G. Wheeler, M. Sc.: "Southern Blue and Fin Whales"-Discovery Reports. Vol. I.
J •.F. G. Wheeler, M. Sc.: "The Age of Fin Whales at Physical Maturity with Notes on Multiple
Ovulations." - Discovery Reports Vol. ll.
Alec H. Laurie, M. A. : "The Age of Female Blue Whales and the Effect of Whaling on the
Stock. Vol." -Discovery Reports Vol. XV·
L. Harrison Matthews, M. A.:"The Sei Wh<'!le, Balaenoptera bolealis" -Discovery Reposts.

Vol. XVlll."
Yoshio Matsuura: Composition of Whales caught in the Antarctic Ocean-Journal of the Japan
Marine Products Society.
Fin whales-

Vol. 8, No. 5.

Blue whales- Vol. 9, No. 1.
Humpback whales- Vol. 9, No. 2.
Japanese Association of Whaling Industries: Biological Survey of Cetaceae in the Northern
Pacific I Whaling Resources. Vol. 9 (1).