A MORPHOLOGICAL NOTE ON THE INTESTINE OF
THE BOUTU WITH EMPHASIS ON ITS LENGTH AND!LEO-COLIC TRANSITION COMPARED WITH
OTHER PLATANISTIDSFUSAO YAMASAKI AND KENJI KITO
Department of Biology, Sapporo Medical College, Sapporo

ABSTRACTThe intestine of the Boutu, Inia geoffrensis (body length 204 cm, female),
was examined macroscopically and light microscopically. The intestine of
the Inia examined was grossly divided into three portions: the small intestine,
the smooth-walled portion, and the colon. The lengths of these parts wereapproximately 4150 cm, 80 cm, and 40 cm, respectively. The total intestinal
length of the lnia, including the 5 cm long rectum and 3 cm long anal canal,
was 4278 cm. Therefore, the intestine was nearly 21 times as long as the
body length. In the interior of the small intestine, except for the proximal
half of the duodenum, a single distinct longitudinal fold involving both themucosa and submucosa could be seen. The smooth-walled portion was thicker
than the small intestine, and had no fold, but its inner surface became fiat andsmooth. The colon, which had several longitudinal folds in its interior, wasthinner than the smooth-walled portion in the initial one-third and thicker
in the remaining portion. Histological differences between the small intestineand the smooth-walled portion and the colon could not be clearly distinguished in our specimen, except for the abundance of goblet cells in the latter
two portions. The ileo-colic transition of the lnia observed was unclear andthis region was compared with other kinds of platanistids.
INTRODUCTION

Morphological investigation of the internal organs, as well as of other body features,is very important in order to clarify the phylogenetic relationship among the fresh
water dolphins of the family Platanistidae, Platanista, Pontoporia, Lipotes, and Inia.
Concerning the morphology of the intestine, there are some observations previously
reported on Platanista (Anderson, 1879; Slijper, 1962/1979; Takahashi and Yamasaki, 1972), Pontoporia (Burmeister, 1869; Yamasaki, Takahashi and Kamiya, 1975),
Lipotes (Chen and Chen, 197 5; Zhou and Li, 1981 ), and Inia (Zhou, Li and Pilleri,1982). However, our morphological knowledge of the Inia's intestine is still insufficient because the investigation by Zhou et al. (1982) treated only Inia boliviensis,
and no detailed description on the intestine of Jnia geojfrensis has so far been reported except for a partial reference to the morphology of the intestine (Yamasaki
et al., 1975). The present paper, as the second report on the morphological study
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of the digestive tract of I. ge~ffrensis (cf. Yamasaki and Kamiya, 1981, on the stomach), deals with the macroscopical characteristics of the intestine, some of whichwere not observed by Zhou et al. (1982) in I. boliviensis, and discusses the intestinal
length and the ileo-colic transition as compared with other platanistids.
MATERIAL AND METHODS

The intestine of the Bou tu (Amazonian dolphin), Inia geoffrensis (body length 204 cm,
female) used for this study was provided by the Kamogawa Sea World, Chiba,
Japan, after death from unidentified disease. The intestine taken from the abdominal cavity was preserved in 10% formalin solution and transported to ourlaboratory. After macroscopical observations, small pieces for the light microscopy were taken from several parts of the intestine, and were embedded in paraffin.
Sections were stained with hematoxylin and eosin.
OBSERVATIONS (PLATE I AND II)

The intestine of I. geojfrensis was observed to be grossly divided into three portions:the small intestine, the colon, and the portion which was located between thesetwo. The intermediate portion was thicker than the small intestine and was alsocharacterized by its smooth interior. It was difficult to determine whether thisportion belonged to the last part of the small intestine or the initial part of thelarge intestine; therefore, for convenience, we called it a "smooth-walled portion"of the intestine in this study.
The initial part of the small intestine, the duodenum, consisted of the duodenal ampulla and the duodenum proper. The former was continuous with thepyloric stomach with a small opening 2 mm in diameter. It was funnel-shaped,
approximately 2.5 cm in diameter at the widest proximal part, about 3 cm in length,
and continued to the duodenum proper. The hepato-pancreatic duct opened in
the duodenal lumen about 9 cm away from the commencement of the duodenum
proper and formed a distinct plica longitudinalis duodeni about 1.5 cm long and0.6 cm wide. Although the border between the duodenum and the jejunum was
unclear, and fairly difficult to determine, the length of the duodenum proper appeared to be roughly 20 cm. The small intestine, except for the proximal half ofthe duodenum, formed a number of coiled loops suspended by a broad mesentery,which was 7-12 cm in width. The small intestine was approximately 0.7-0.8 cm
in diameter but in several parts there was a slight dilation, about 1 cm in diameter,
extending over a length of about 5-10 cm. The length of the small intestine, fromthe initial part of the duodenum to the beginning of the smooth-walled portion,measured nearly 4,150 cm. No circular folds could be seen in the interior of any
parts of the small intestine. The inner surface of the duodenal ampulla exhibitedseveral folds, which might be not present in the living state, being slightly irregular;
and at about 5 cm from the opening of the hepato-pancreatic duct one single distinct
longitudinal fold began and continued throughout almost the whole interior of theSci. Rep. Whales Res. Inst.,
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167

small intestine (Fig. 1). This single fold projected into the intestinal lumen beyond a half diameter of the lumen and was generally found on the antimesentericborder (Fig. 6). The fold became slightly smaller and irregular in the several
dilated parts of the small intestine mentioned above, and also near the smoothwalled portion (Fig. 2). The smooth-walled portion of the intestine measured
about 80 cm long. This portion, about 1 cm or more in diameter, was clearly
thicker than the small intestine. The longitudinal folds could not be seen in thisportion and the inner surface became flat and smooth (Fig. 3). However, from
one to three short longitudinal folds, 2-5 cm long, were seen in some places of thisportion. The smooth-walled portion formed an intestinal loop but its mesentery
was shorter than in the small intestine. The vessels in the mesentery made an
acade and many vasa recta reached the smooth-walled portion as seen in the small
intestine, though this pattern tended to become irregular in the distal part. The
last part of this portion ran upwards and flexed dorsal, and continued to the nextcolon.
The colon descended straightly and continued to the rectum. The mesenteryof the colon was short, about 5-6 cm wide, and gradually reduced in width analwards, disappearing in the anal part. The initial one-third of the colon was somewhat thinner than the smooth-walled portion, with a width of approximately 1 cm
or less, and the last two-thirds became thicker, being about 1.5 cm in diameter. In
the interior of the colon there could be seen from two to five distinct folds runninglongitudinally (Fig. 4 ), which became more marked in the last two-thirds (Fig. 5 ).
These folds were broader than those in the small intestine. Most folds continued
throughout the entire length of the colon, but short ones, 2-3 cm long, were present
in some places. The colon was approximately 40 cm long, and the rectum andthe anal canal were 5 cm and 3 cm long, respectively.
The thickness of the mucosa of the duodenal ampulla was 0.6-0.7 mm, andthat of the rest was slightly thinner, being 0.4-0.5 mm in thickness. Since thespecimen observed was in poor condition for histological examinations, typical villi
could not be clearly observed in the small intestine (Fig. 7) and it could not be
determined whether villi were present or not in the smooth-walled portion (Fig. 8).
Intestinal glands were mucous in the duodenal ampulla. Those of the rest of theintestine morphologically resembled each other. But the glands of the smoothwalled portion and colon showed some tertuous courses and goblet cells became
abundant, especially in the bottom of the glands (Figs 8-10). Musclaris of thewhole intestine, consisting of thick inner circular and thin outer longitudinal layers,
was 0.3-0.5 mm thick in the small intestine (Fig. 6) and smooth-walled portion,and 0.5-0.8 mm thick in the colon. The inner muscular layer was 4-5 times as
thick as the outer one. The sphincteric structure could not be seen at either theborder between the small intestine and the smooth-walled portion or between thisportion and the colon. The folds observed in the whole intestinal tract involved
both the mucosa and the submucosa (Fig. 6). Some accumulations of lymphoidtissue were found in the lamina propria of the colon but well-developed lymphatic
nodules could not be seen in any portion of the intestinal tract of the observed
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YAMASAKI AND KITO

specimen.
DISCUSSION

The total length of the whole intestine of the I. geojfrensis (body length 204 cm) was
4,278 cm; thus the ratio of the intestinal length to the body length is nearly 21 to 1.
This agrees well with the ratios, 19 and 20 to 1, obtained in I. boliviensis (bodylengths 145 cm and 174 cm, respectively, Zhou et al., 1982). The ratios in otherfresh water dolphins reported by previous investigators are shown in Table 1 *.
Platanista's ratio is notably low and Pontoporia's is higher by far than that of otherdolphins. Since there appears to be a tendency for the intestinal length to increase
with age in our collected specimens of Platanista and Pontoporia (cf. Takahashi andYamasaki, 1972; Yamasaki et al., 1975), the ratio becomes slightly higher when
specimens are limited to adults.
TABLE I.
SpeciesInia
bo!iviensis
geoffrensis
Platanista
gangetica

THE RATIO OF THE INTESTINAL LENGTH TO THE
BODY LENGTH IN PLATANISTIDS
Ratio

19.0-20.0
21.0

Zhou et al. (1982)
Yamasaki et al. (1975) and present study (1983)

4.1-4.4
7.3

Anderson (1879)
Slijper (1962/1979), no descriptions on the length of bodyand intestine
Takahashi & Yamasaki (1972)

3.6-4.9

Pontoporia
blainvillei

Lipotes
vexillifer

Authors and Remarks

32.0
24.0-37.0

Burmeister (1869)
Yamasaki et al. (1975), examined in unfixed state in most
cases

15.0

Chen & Chen (1975), ratio of whole length of the digestiv~ tract, from mouth to anus, to the body length
Zhou & Li (1981)

12.5-14.9

In the small intestine of the Inia observed there are no circular folds, but asingle distinct longitudinal fold projecting into its lumen can be seen as in Pontoporia (Yamasaki et al., 1975). Although Zhou et al. (1982) did not mention thisfold in I. boliviensis, the fold in I. geojfrensis is plainly visible as a permanent structure and can also be easily felt through the wall of the intestine when the intestineis grasped between the fingers. In Platanista, well-developed circular folds are present on the inner surface of the small intestine but the surface of about the last
50 cm of the ileum becomes fiat and smooth, and a distinct longitudinal fold, about
30 cm long, is present at this portion (Takahashi and Yamasaki, 1972). The presence of the circular or longitudinal folds in the small intestine is due to the struc-* The ratios of Platanista, Pontoporia, and Inia misquoted in our previous paper (1981) are corrected as shownin Table 1.

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169

tural specializations that serve to increase the area of the surface exposed to thelumen. It is assumed that numerous well-developed circular folds are a much
more effective means of augmenting the surface area of the mucosa than a single
longitudinal fold. This fact may have some relation to the short intestine of Platanista and to the long intestine of lnia and Pontoporia. In the reports of the intestine of Lipotes (Chen and Chen, 1975; Zhou and Li, 1981) there are no macroscopical descriptions of its internal structure, and therefore it is unclear whether
Lipotes' intestine has such longitudinal folds or not.
In cetaceans it is generally known that there is no clear external transition between the small and the large intestines; except for Platanista and mysticetes whichhave a short caecum. In Pontoporia the external appearance of the transition between the small and the large intestines was not so difficult to determine because of
the differences in thickness, the supply of the mesenteric vessels, and the interior
equipped with a single longitudinal fold in the small intestine while there were two
folds in the large intestine (Yamasaki et al., 197 5 ). However, in the case of ourInia, it was very difficult to determine the transition between the small and thelarge intestines. In the smooth-walled portion the single longitudinal fold disappears and the inner surface becomes flat and smooth. Nevertheless, the pattern ofthe blood supply, the vasa recta, seen in the small intestine was observed until the
distal part of the smooth-walled portion. Since our specimen was, unfortunately,in poor condition for histological examination, neither the presence of typical villi,
which is one of the most important characteristic features of the small intestine,
nor the disappearance of the villi in the large intestine could be observed. In addition, other histological findings generally found in each small and large intestinein mammals in general could not be observed, so that we did not confirm thetransition region from the small to the large intestine. However, goblet cells of
the intestinal glands of our lnia were abundantly observed in the smooth-walled
portion and the colon, unlike in the small intestine. This distribution of the goblet
cells in the smooth-walled portion and colon in lnia appears to correspond well tothat in the ascending and descending colons in Pontoporia.
Zhou et al. (1982) did not mention this smooth-walled portion, and they considered the colon to be only the portion, about 35 cm long, which ran caudally
connecting with the rectum, and they described the presence of villi in the small
intestine. The length of the large intestine of Pontoporia was 80 cm or more (Yamasaki et al., 1975) and in Lipotes it is about 100 cm (Zhou and Li, 1981). As the
structure of the intestine of our lnia resembles that of Pontoporia, and probably
Lipotes, rather than that of Platanista, the 35 cm long colon of I. boliviensis seems tobe proportionally too short. Although in our previous paper (Yamasaki et al., 1975)
the smooth-walled portion was considered as ascending colon, there are no conclusive characteristics to determine whether the smooth-walled portion belongs to
the small or the large intestines, as is discussed in this study. In order to clarify
the subject of the ileo-colic transition, it is necessary to investigate cetacean species
having no caecum from comparative morphological viewpoints.
Zhou et al. (1982) observed no goblet cells in the intestinal glands in I. boliviSci. Rep. Whales Res. Inst.,
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YAMASAKI AND KITO

ensis and also stated that this is one of the characters common to Inia and Platanista.
We found abundant goblet cells in /. geojfrensis, as in Pontoporia and Lipotes (cf.
Yamasaki et al., 1975; Zhou and Li, 1981). Though present, the cells were muchless abundant in Platanista (Takahashi and Yamasaki, 1972). Therefore, the absence of the goblet cells may be peculiar to I. boliviensis among the fresh water dolphins previously reported.
Although well-developed lymphatic nodules in the lamina propria were present from the last part of the ileum to the initial part of the colon in Platanista(Takahashi and Yamasaki, 1972) and in Pontoporia (Yamasaki et al., 197 5 ), development of lymphatic tissues of the intestinal tract was very poor in our Inia. Zhou
et al. (1982) described numerous lymphatic nodules aggregating from the last partof the ileum (probably corresponding to the last part of the smooth-walled portionin our case) to the anal part, and well-developed anal tonsils. Since the anal
tonsils in our Inia were poorly developed (Yamasaki, Komatsu and Kamiya, 1977),
poorly developed lymphatic tissue in the intestinal tract seems due to individual
characteristics; well-developed lymphatic nodules could not be seen anywhere in
the intestinal tract.
Pilleri and Gihr (1977, 1980) and Pilleri, Marcussi and Pilleri (1982) statedthat there were clear morphological and morphometric differences between the
Bolivian and Amazonian Inia, based on the observations of their brains and teeth.
The stomachs of both I. geojfrensis and /. boliviensis quite resemble each other instructure (cf. Yamasaki and Kamiya, 1981; Zhou et al., 1982). However, as mentioned above, there are some remarkable macroscopical differences in their intestines. It remains a problem whether such differences in the intestine were interspecific or not, because the present observations on Inia were based on one case.
Therefore, further observations of this subject, based on the sufficient materials,
should be done in order to compare with each other.
ACKNOWLEDGEMENTS

We wish to thank Dr T. Tobayama, Kamogawa Sea World, Chiba, for kindly supplying the specimen for this study.
REFERENCES
ANDERSON, J., 1879. Anatomical and zoological researches: comprising an account of the zoological results of the two
expeditions to Western Yunnan 1869 and 1875; and a monograph of the two cetacean genera, Platanista and Orcella.
2 vols. Qurritch, London.
BURMEISTER, H., 1869. Pontoporia blainvillii Gray. In: Descripcion de cuatro especies de de!finides de la costa
Argentina en el Ocifano Atltintico. Anal. Mus. Publico Buenos Aires, 1: 384---445.
CHEN, I. and W. CHEN, 1975. Notes on some morphological and anatomical features of the white-flag dolphin, Lipotes vexillifer. Acta Hydrobiol. Sinica, 5: 360-368. (in Chinese with English summary).
PrLLERI, G. and M. GrnR, 1977. Observations on the Bolivian (Inia boliviensis d'ORBIGNY, 1834) and the
Amazonian Bufeo (Inia geoffrensis de BLAINVILLE, 1817) with description of a new subspecies (Iniage~ffrensis humboldtiana). Investigations on cetacea 8: 11-76.
PrLLERI, G. and M. GrnR, 1980. Checklist of the cetacean genera Platanista, Inia, Lipotes, Pontoporia, Sousa
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and Neophocaena. Investigations on cetacea, 11: 33-36.
P1LLERI, G., G. MARCUZZI and 0. PILLER!, 1982. Speciation in the Platanistoidea. Systematic, zoogeographical and ecological observations on recent species. Investigations on cetacea, 14: 15-46.
SLIJPER, E.J., 1962/1979. Whales. Hutchinson, London. (English translation).
TAKAHASH., K. and F. YAMASAKI, 1972. Digestive tract of Ganges dolphin, Platanista gangetica. II. small
and large intestines. Okajimas Fol. anat.jap., 48: 427-452.
YAMASAKI, F., K. TAKAHASHI and T. KAMIYA, 1975. Digestive tract of La Plata dolphin, Pontoporia blainvillei. II. small and large intestines. Okajimas Fol. anat.jap., 52: 1-26.
YAMASAKI, F., S. KOMATSU and T. KAMIYA, 1977. A comparative morphology of anal tonsils in Platanistidae. Sci. Rep. Whales Res. Inst., 29: 95-100.
YAMASAKI, F. and T. KAMIYA, 1981. The stomach of the Boutu, Inia geoffrensis: comparison with those ofother platanistids. Sci. Rep. Whales Res. Inst., 33: 69-81.
ZHou, K. and Y. LI, 1981. On the intestine of the Baiji, Lipotes vexillifer. Acta :(,ool. Sinica, 27: 248-253. (inChinese with English summary).
ZHou, K., Y. LI and G. PILLER!, 1982. The digestive tract of Inia boliviensis (D'ORBIGNY, 1834). Investigations on cetacea, 13: 125-133.

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EXPLANATION OF PLATESPLATES I AND II
Inner views of several parts of the intestine of Inia geoffrensis (1-5). The intestine is
opened wide to show its interior at the mesenteric border. Figs 1-5, actual size.
Fig. 1. Inner surface of the ileum at about 5 m proximal to the smooth-walled portion. Note that a single distinct longitudinal fold can be seen and no circularfolds are present in the small intestine.
Fig. 2. The transitional part between the small intestine (left) and the smooth-walled
portion (right). The longitudinal fold becomes slightly irregular in the last part ofthe small intestine and the inner surface becomes smooth in the smooth-walled
portion.
Fig. 3. Mid-part of the smooth-walled portion, which is thicker than the small intestine. Short longitudinal folds are seen in some places in this portion.
Fig. 4. The last part of the smooth-walled portion (left) and the initial part of thecolon (right). Two or three longitudinal folds are present in the initial part of thecolon.
Fig. 5. Mid-part of the colon. Marked longitudinal folds can be seen in the surface.
Fig. 6. Cross section of the mid-part of the small intestine. A prominent longitudinalfold, involving both the mucosa and the submucosa, projects into the lumen fromnear the antimesenteric borders. x 11.
Fig. 7-10. Same magnification photomicrographs of the mucosa of the several partsof the intestinal tract. x 10. Mid-part of the small intestine (Fig. 7), mid-part ofthe smooth-walled portion (Fig. 8), initial part of the colon (Fig. 9), and mid-partof the colon (Fig. 10). Typical villi could not be seen in the small intestine in our
specimen. The intestinal glands of the small and the large intestines are rather
straight. Goblet cells become abundant in the smooth-walled portion and thecolon.

Sci. Rep. Whales Res. Inst.,

No. 35, 1984

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