NationalLibraryOfScotland/medical-history-of-british-india huggingface dataset:
No. 30.
(NEW SERIES. )
SCIENTIFIC MEMOIRS
BY
OFFICERS OF THE MEDICAL AND SANITARY DEPARTMENTS
OF THE
GOVERNMENT OF INDIA.
THE THEORY AND PRACTICE OF ANTI-RABIC
IMMUNISATION.
BY
CAPTAIN W. F. HARVEY M.B., I.M.S.
AND
CAPTAIN ANDERSON MCKENDRICK, M.B., I.M.S.
ISSUED UNDER THE AUTHORITY OF THE GOVERNMENT OF INDIA
BY THE SANITARY COMMISSIONER WITH THE GOVERNMENT
OF INDIA, SIMLA.
CALCUTTA
SUPERINTENDENT OF GOVERNMENT PRINTING, INDIA
1907
P r i c e A n n a s 1 2
or 1s. 2d.
NCAR—VSI 4/34.
Vol. IV
1984
THE
Indian Journal of Veterinary Science
AND
Animal Husbandry
Issued under the authority
of
The Imperial Council of Agricultural Research.
Published for
The Imperial Council of Agricultural Research
BY
MANAGER OF PUBLICATIONS, DELHI.
PRINTED BY THE MANAGER, GOVERNMENT OF INDIA PRESS, NEW DELHI
1936.
Price annas 3 or 4d.
ORIGINAL ARTICLES
A COMPARATIVE STUDY OF SCHISTOSOMA
SPINDALIS,
MONTGOMERY, 1906 AND SCHISTOSOMA NASALIS, N. SP.
BY
M. ANANT NARAYAN RAO,
Lecturer in Parasitology, Madras Veterinary College.
(Received for publication on 27th October 1933.)
(With Plates I-V and five text-figures.)
CONTENTS.
INTRODUCTION . . . . . . . .
OVA . . . . . . . .
MIRACIDIA
.
.
.
.
.
.
.
.
INTERMEDIARY HOST . . . . . . . .
CERCARIAE . . . . . . . .
EXPERIMENTAL INFECTION AND INCUBATION PERIOD . . . . . . . .
EARLY SYMPTOMS IN EXPERIMENTAL CALVES INFECTED WITH CERCARIA
INDICAE XXX
.
.
.
.
.
.
.
.
POST-MORTEM FINDINGS . . . . . . . .
SOME OBSERVATIONS IN HISTOPATHOLOGY . . . . . . . .
EXPERIMENTAL INFECTION WITH CERCARIAE OF S. SPINDALIS . .
POST-MORTEM FINDINGS IN CALVES INFECTED WITH S. SPINDALIS . .
SOME OBSERVATIONS ON SCHISTOSOMES COLLECTED FROM ANIMALS IN
THE ABATTOIRS
.
.
.
.
.
.
.
.
THE HABITAT OF S. SPINDALIS AND THE NASAL SCHISTOSOME . . .
DISCUSSION ON SOME POINTS OF DIFFERENCE IN THE MORPHOLOGY OF
S. SPINDALIS
AND THE NASAL SCHISTOSOME
.
.
.
.
.
.
.
.
THE IDENTITY OF THE NASAL SCHISTOSOME . . . . . . . .
SPECIFIC DIAGNOSIS . . . . . . . .
GENERAL REMARKS . . . . . . . .
ACKNOWLEDGMENTS . . . . . . . .
REFERENCES . . . . . . . .
PAGE.
l
3
6
9
10
12
13
14
15
17
18
19
19
21
24
24
24
27
27
INTRODUCTION.
In the domesticated animals in India four species of Schistosomes are known to
exist, one being S. bovis Sonsino, 1876, and the other three being S. bomfordi,
S. spindalis and S. indicum discovered by Montgomery in 1906. All these Schisto-
B 2
2
THE INDIAN JOURNAL OF VETERINARY SCIENCE AND ANIMAL HUSBANDRY [ IV, I.
somes are found in the portal veins or mesenteric veins or both, of cattle, sheep,
goats, buffaloes and horses, as the case may be. In the Madras Presidency the
author has met with only two species hitherto, viz., S. spindalis and S. indicum in
the portal system of those animals and, so far as he is aware, there appears to be no
record of any other species having been found excepting the nasal schistosome
recently discovered in the veins of the nasal fossae of cattle.
Twelve years after Montgomery [ 1906 ] published his paper on those three
Schistosomes discovered by him, the life-history of one—S. spindalis—was worked
out by Liston and Soparkar [ 1918 ]. Almost another decade elapsed before Fairley
and Mackie (1925) studied the histopathology of S. spindalis in goats and proved
that it affects exclusively the portal system and its ova are voided only with the
faeces. Liston and Soparkar [ 1918 ] also found that the ova of S. spindalis pass
out in faeces of the infected animal. The other two Schistosomes of cattle dis-
covered by Montgomery have so far been found to develop only in the portal
system; consequently their ova are found in the faeces of the host. As regards
S. bovis, it has been proved by MacHattie and Chadwick [ 1932 ] and others that
it chiefly inhabits the portal system and the ova of these find an outlet only through
the faeces of the host. In man, two out of the three species, develop in the portal
system and the only exception being S. haematobium (Weinland 1858) which deve-
lops usually in the pelvic plexus and its ova are voided chiefly in the urine. It is
interesting to note that in nasal schistosomiasis in cattle the species of Schistosome
concerned chooses the nasal veins as its habitat and its ova are found only in the
nasal discharge, a departure analogous to that of S. haematobium in man. In these
two species, therefore, the site of infection in their normal hosts is different from
that of the other species found in man or cattle.
Sometimes two or more species of Schistosomes are found in the same host.
Montgomery [ 1906 ], for example, found S. bomfordi and S. spindalis in the same
animal and he was able to distinguish the one from the other by noting the difference
in the morphology of the worms and by the shape of their ova. Leiper [ 1915 ]
writes—" A comparative study of mature specimens of S. bovis and S. haematobium
showed that those two species were so closely allied that when experimentally reared
in an abnormal host, a differential diagnosis could only be made with certainty upon
the characters of the fully grown worms and especially by those of the egg shell'.
The last remark of Leiper is further strengthened by MacHattie and Chadwick
[ 1932 ], who say that " the eggs of the various Schistosomes from man and animals,
within certain limits of variation, have a definite shape which is generally regarded
as being characteristic of the species ". It would appear, therefore, that the mor-
phology, the characteristic shapes of the ova and the site of infection are important
SCHISTOSOMA SPINDALIS, MONTGOMERY, 1906 & SCHISTOSOMA NASALIS, N. SP. 3
points in arriving at an identification of the species of Schistosomes. In addition to
this, the morphology of the miracidium, and cercariae should also help in the
diagnosis, where the life-history of the worm is known.
Bhalerao [ 1932 ], in the course of his paper observes that:—
(1) much importance cannot be attached to the question of the number
of testes present in the male schistosome.
(2) the length of the common caecum cannot be taken into account
owing to its variable nature.
(3) the dimensions of the body cannot be given any importance since
they vary with age.
(4) the tuberculate or non-tuberculate nature of the cuticle should not
be considered as of any value in distinguishing one species of schis-
tosome from another.
(5) polymorphism in eggs appears to be the rule in many species of
schistosomes.
(6) the site of infection does not help matters.
It would appear therefore that the specific diagnosis of a schistosome has no
value, when it is based on the above points. Hence Bhalerao's diagnosis of the
Nasal Schistosome as no other than S. spindalis Montgomery [ 1906 ] cannot be
accepted in the light of his own observations, since he arrives at a diagnosis by
comparing, in a tabular form, some of the features enumerated above. Three other
workers, however, consider that the nasal schistosome is different from S. spindalis.
Datta [ 1932 ], says that the parasite resembles S. spindalis, but certain differences
suggested to him that it may be a new species. Malkani [ 1933 ], thinks that it is
a variety of S. spindalis. Rao [ 1932 ], describes it as a new species. Now that
the life-history of the nasal schistosome has been worked out by Rao [ 1933 ] and
that of S. spindalis has already beed worked out by Liston and Soparkar [ 1918 ], it
would be interesting to compare these species from different standpoints, such as
the shape of the egg, morphology of the miracidium, cercaria and of the worms to
arrive at a specific diagnosis.
THE OVA.
The author had opportunities to study the ova of Nasal Schistosomes in the
nasal washings obtained from over 300 cases from different parts of the Presidency
4
THE INDIAN JOURNAL OF VETERINARY SCIENCE AND ANIMAL HUSBANDRY [IV, I.
and also from three experimentally infected calves. The shape of the ovum is an
elongated spindle. The middle third of the body of the ovum, which is concavo-
convex, is prolonged into a horn on either side, one of which has its extremity
bluntly rounded and the other terminates in a spine. The spine is straight in some
ova, but in the majority of mature ova, it is curved slightly or sharply like a hook,
towards the concave margin of the body. In none of them, however, was the spine
seen curved towards the convex margin of the body of the egg. The horns them-
selves are either straight or slightly turned up towards the convex margin of the
egg. With proper orientation the typical shape of this " zygomorphic " ovum is
always seen. In the fresh nasal discharge, the mature ovum shows a miracidium
inside it and it appears that in the majority of the ova the snout of the miracidium
is pointed towards the spine end of the ovum. In the egg, the miracidium occupies
only the central area and the horns of the egg seem to contain a clear fluid. This
fluid in the horns of the egg becomes apparent when some tap water is added to the
fresh nasal washings. A seething mass of minute bubbles arise in this fluid round
about the miracidium. (Pl. III, fig. 4.) Presumably, an increase of these
bubbles brings about a rupture of the egg shell, liberating the embryo. This
confirms Malkani's [1933] observations regarding this phenomenon.
The ovum of S. spindalis is also spindle-shaped, zygomorphic and has a body
and two horns. In this ovum there is a bulge on one side of its body and the
opposite side is straight throughout its length or even slightly convex at its middle.
The spine is straight or curved towards the straight border and in over 15 per cent.
of these ova, the spines are curved upwards towards the convex side of the body
of the egg. The horns are usually straight and in a few ova they are slightly
inclined towards the straight side and they are never seen to bend towards the
bulged side of the egg as in the ova of nasal schistosomes.
Datta [1932] has published a photograph of a typical ovum from the nasal
washings and in that the bend of the horns towards the convex side of the body
is clearly seen. Malkani [1933] says that the ovum found in the nasal washings has
a concave border in place of the straight border found in the ova of S. spindalis.
He has also given a photograph showing the typical shape of the ova. Rao [1932]
has published a photograph of the ovum of S. spindalis and another of the nasal
schistosome to show the difference in their shape and size. Bhalerao [1932] has
illustrated his paper with drawings of ova in nasal washings and admits that they
usually possess, more or less, a degree of curvature on one side. He also says that
the egg of nasal schistosome presents different appearances when viewed in different
positions, and this is to be expected in a zygomorphic ovum. This is also true
with the ovum of S. spindalis.
PLATE I.
1. Ovum of S. spindalis. Note the shape and actino-body round the
horn behind the spine.
2. Ovum of S. nasalis in uterus. Note shape. × 60. Photograph of
fresh material.
3. Ova of S. spindalis in uterus. Note shape. × 60. Photograph of
fresh material.
4. Section of liver of experimentally infected snail showing cercariae
in sporocysts.
SCHISTOSOMA SPINDALIS, MONTGOMERY, 1906 & SCHISTOSOMA NASALIS, N. SP.
6
Montgomery [1906], Tanabe [1925], Baylis [1929] and Faust [1930] have
shown, in their drawings of Schistosome ova, that one side of the ovum of
S. spindalis is flat. Fairley and Mackie [1930] have given a diagram of a female
S. spindalis with ova in its uterus and also of a few mature ova. These illustra-
tions show that one side of the ovum is flat or even slightly convex and the other
side is bulged in the middle. In some of these ova the spines are curved towards
the bulged border. Photo-micrographs (Pl. I, figs. 2 & 3) of fresh female
schistosomes show the difference in shape and size of the ova in utero. Thus there
appear to be differences in the shape of the ova of these two species, and these are
tabulated below for comparison.
TABLE I.
—
Length* . . .
Breadth . . .
Spine
.
.
.
.
Horns
.
.
.
.
Base*
.
.
.
.
Where found . . .
Ova of S. spindalis.
284—400
80—90
Often curved towards the bulged
border.
Straight or slightly curved towards
the flat side.
Flat or slightly convex in its
middle.
Only in faeces . . .
Ova of nasal schistosome.
336—581
60—80
Often curved towards the concave
border.
Straight or slightly curved towards
the bulged border.
Concave.
Only in the nasal discharge.
* The length and breadth are in microns and include the maximum and minimum given by various
authors. The base represents the side opposite to the bulged margin of the body of the egg.
Datta [1932] says that the type of ova met with in the nasal discharge has not
been found in the faeces or urine of animals having nasal schistosomiasis.
Malkani [1933] confirms this. The author examined samples of faeces and urine
of 86 cases, and in addition he has frequently examined the faeces and urine of the
three experimentally infected calves and agrees with Datta and Malkani. Further
the author did not find ova typical of S. spindalis in the nasal washings. Therefore
even in the same animal, it would appear that the two species of Schistosomes
with the two different types of ova, are met with in two different situations
respectively. From the table of differences it is evident that the ovum of the nasal
schistosome is different from the ovum of S. spindalis.
6
THE INDIAN JOURNAL OF VETERINARY SCIENCE AND ANIMAL HUSBANDRY [ IV, I.
Malkani [1933] says that he found in the nasal washings only one type of egg
with one typical shape thus confirming Rao's [1932] finding. Since then the
author has examined a large number of specimens of nasal discharge, preserved in
3 per cent. NaOH or KOH solution, received from the districts and also fresh
nasal washings from the three experimentally infected calves. None of the ova
showed any departure from the typical shape described above. Occasionally some
immature ova showed a narrower contour and a smaller size than the rest. Even
in the sections of tissue the typical shape of the ova is seen provided the ovum is
cut longitudinally about its middle. Datta[1932], however, met with a few ova
having peculiar shapes, with or without spines or even having two spines one on
either end. These ova have been illustrated by Bhalerao [1932]. These shapes do
not appear to be within reasonable limits of variation in a given species and await
confirmation.
Bhalerao [1932] suggested that the central bulge on one side of the ovum of
nasal schistosome is probably due to some organ of the developing embryo. The
present author has verified this and found that the bulge in the body of the ovum
of either the nasal schistosome or of S. spindalis is not caused by the developing
miracidium or any of its organs. The centrally placed miracidium in an egg has
failed to show any protrusion of its part or parts beyond its usual almond-shaped
contour during its development and apparently the miracidium is only a passive
entity within the characteristically shaped shell moulded in the ootype. It has also
been seen that even the active movements of a miracidium inside the egg, have no
effect whatever even to temporarily alter the shape of the rigid egg shell.
THE MIRACIDIA.
In the fresh nasal discharge, particularly in that from recently infected animals,
fully developed miracidia are seen in the majority of the ova. Often free miracidia
are encountered in the undiluted discharge particularly when the discharge is thin or
if it is kept in a tube for some little time before examination. This confirms
Malkani's [1933] finding of free miracidia in fresh nasal discharges.
The mature ova of S. spindalis obtained in the rectal scrapings, on the other
hand, do not seem to rupture unless water is added to the sample, when bubbles
develop inside the ovum, just as in the case of nasal schistosome ova, before the
shell bursts.
The miracidium of the nasal schistosome, on emerging from the egg, swims
about vigorously in water with the aid of cilia present on its cuticle. It appears to
SCHISTOSOMA SPINDALIS, MONTGOMERY, 1906 & SCHISTOSOMA NASALIS, N. SP.
7
be positively phototrophic and can live in tap water up to about 24 to 36 hours at
laboratory temperature. The shape of the miracidium is more or less like that of an
almond with its shoulders slightly broadened at the anterior extremity, and it
appears to be slightly flattened dorsoventrally. The anterior extremity bears on it,
a protrusible semi-circular snout without any cilia. The posterior extremity is
bluntly rounded and is covered over with cilia as on the other parts of the cuticle.
Cilia are absent round two papillae, one on each shoulder, and each of these papillae
is the common opening of the ducts of a cluster of small secretory glands which will
be referred to later. About the middle of the body of the miracidium is a girdle of
groups of cilia running right round the body. A little behind this girdle, around
each of the two excretory pores, cilia are absent. The girdle of groups of cilia is an
interesting morphological detail. This is seen when the surface of the cuticle is
carefully brought into focus of the microscope and it is best seen when the ciliary
movements are slowing down. In the miracidium of the nasal Schistosome the
girdle is formed by 10 to 12 groups of cilia across the visual section, and each group
is cone shaped.
On the semi-circular snout is the small oral opening which leads into a short
tubular or saccular gut containing granular material. The gut extends backwards
along the median line, to a rounded bunch of small secretory glands or may even
slightly overlap this organ. On either side of the gut is a long and fairly large
secretory gland, containing large oxyphilic granules, the duct of each opening on the
postero-lateral margin of the snout. These glands, which may be called the anterior
or lateral secretory glands, are very much longer than the gut. Behind the gut,
in the median line is a fairly large circular looking object consisting of a group of
small secretory cells or glands showing fine basophilic granules. This group appears
to be a pair placed close together. The ducts of each pair of groups open on a
peg-like papilla on each shoulder of the miracidium. Surrounding these two groups
of secretory cells is the nerve tissue with a few short linear prolongations proceeding
posteriorly into the parenchyma of the miracidium. In the body of the miraci-
dium, particularly in the posterior region, there is a fairly large number of rounded
germinal cells. These germinal cells, apparently, are derived from the germinal
epithelium lining the inside of the cuticle.
The excretory system consists of an anterior and posterior flame cell in each
mesial half of the body. One flame cell is situated just near the rounded group of
small secretory cells and from it a collecting tubule arises to join a similar tubule,
which takes its origin from the posterior flame cell situated near the excretory pore.
Each of the common tubes formed by the junction of these two tubules, at first
passes forwards, re-curves on itself, proceeds backwards and at about the middle of
8
THE INDIAN JOURNAL OF VETERINARY SCIENCE AND ANIMAL HUSBANDRY [ IV, I.
the body it forms a tangle and ultimately opens in the excretory pore, which is at
the margin of the commencement of the posterior third of the miracidium.
(Text-fig. 1.)
FIG. 1. Miracidium Schistosoma nasalis.
The miracidium of S. spindalis has not been fully described hitherto and this
opportunity is taken to do so to facilitate comparison. The miracidium of S.
spindalis is also almond shaped, has a ciliated coat and appears to be smaller in size
than the other one. The snout is smaller and the oral opening leads into a tubular
gut containing granules as in the other. On either side of this gut is a short saccular
anterior secretory gland, shorter than the gut and contains oxyphilic granules. The
duct of each of these glands opens in the same situation as in the other. Behind the
gut in the median line the two groups of secretory cells are exactly as in the other
miracidium and their ducts open on the two peg-like papillae as in that one. The
arrangement of the nerve tissue and the excretory system is similar to that of the
other miracidium but the common excretory tubes do not proceed forwards to such
an extent and are less convoluted. There are breaks in the ciliary coat only around
the secretory and excretory pores and a girdle of ciliary groups is present. It is
interesting to note that the number of groups of cilia on this girdle varies from 6 to
8 across the visual section. The body contains larger and more or less quadrangular
SCHISTOSOMA SPINDALIS, MONTGOMERY, 1906 & SCHISTOSOMA NASAL1S, N. SP. 9
germinal cells when compared to the other. (Text-fig. 2). The. differences between
FIG. 2. Miracidium of Schistosoma spindalis.
S 1. } Secretory glands.
S. 2.
F. O. Flame cell.
E. P. Excretory pore.
G. E. Germinal
epithelium.
the two miracidia are summed up in the following table: —
TABLE II.
1. Anterior excretory glands .
2. The gut
.
.
.
.
3. No. of ciliary groups on the
girdle .
.
.
.
.
4. Germinal cells . . .
Miracidium of S. spindalis.
Shorter than gut
.
.
.
.
Longer than the anterior secretory
glands .
.
.
.
.
. .
6 to 8
. . . . . .
Quadrangular and large . . .
Miracidium of
the nasal
schistosome.
Longer than gut.
Shorter than the
anterior secretory
glands.
10 to 12.
Rounded and small.
THE INTERMEDIARY HOST.
Rao [1933] successfully infected three calves with Cercariae Indicae XXX
Sewell 1922, which be obtained from two species of snails, viz., Limnea leuteola (Lam-
10 THE INDIAN JOURNAL OF VETERINARY SCIENCE AND ANIMAL HUSBANDRY [ IV, I.
arck) and Planorbis exustus (Deshayes). He observed that the infection rate was
more in Limnea species than in Planorbis species. Sewell [1922] found that Cer-
cariae Indicae XXX predominated in Limnea accuminata than in Planorbis exustus.
As regards the cercariae of S. spindalis, Soparkar [1921] has stated that these were
always found in Planorbis exustus and very rarely in Limnea accuminata. Fairley
and Mackie [1930] obtained cercariae of S. spindalis only from Planorbis exustus in
Bombay for their work. If this has any significance, then it shows that the Nasal
Schistosome prefers Limnea species as its intermediate host and S. spindalis chooses
Planorbis species. It is evident, therefore, that the Nasal Schistosome is less selec-
tive in choosing its intermediate host than S. spindalis.
THE CERCARIA.
The cercaria concerned is small, apharyngeal, and brevifurcate. It is a feeble
swimmer and in a test tube of water its movements are mostly up and down. It
swims with its tail foremost and is positively phototrophic. The body surface is
covered over with small thin spines pointing backwards. The tail stem and furcal¬
rami are covered over with coarser spines comparatively and these are curved
slightly at their ends.
The body of the cercaria is pearshaped and on its wider end the forked tail is
inserted. In the anterior end of the body is a muscular and oval penetrating organ
crowned with ten hollow spines, five on each side of the median line. The ventral
sucker is situated at the commencement of the posterior fourth of the body and is
lined with three or four rows of fine spines. When the ventral sucker is held con-
tracted, it usually shows a " Y " shaped depression on it.
The alimentary system consists of an oral opening situated subterminally on the
penetrating organ, leading into a fine tubular œsophagus which ends in a dilated sac
in front of the ventral sucker.
The secretory system consists of five pairs of secretory glands arranged on
either side of the ventral sucker and in the space between it and the posterior end of
the body. On each side, the anterior pair of glands are filled with coarse granules
and the posterior three pairs contain fine granules. Their ducts open individually in
the pointed hollow spines referred to already.
The genital system is represented by a group of cells immediately behind the
ventral sucker. A few cells in the posterior and lateral margins of the body are seen
and apparently these are parenchymatous cells and not rudiments of testes as Sewell
[1922] considers.
The excretory system is composed of a small round bladder situated at the
junction of the body and the tail. From the bladder two canals proceed forwards,