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 Another Random Document on
Scribd Without Any Related Topics
The males have bodies that widen out
considerably behind the ventral sucker, the
lateral parts of which in-roll ventrally, forming
the almost completely closed canalis
gynæcophorus, within which the female is
enclosed. There is no cirrus pouch. The male
has five or six testes, the females are
filiform; the uterus is long. There is no
Laurer’s canal. The ova almost equally
attenuated at either extremity; they have a
small terminal spine, and are not provided
with a lid. They contain a miracidium, ciliated
on all sides, which is characterized by the
possession of two large glandular cells, which
discharge anteriorly beside the gastric sac.
They live in the vascular system of mammals.
(An allied genus [Bilharziella] lives in the
blood-vessels of birds.)

Schistosoma hæmatobium,
Bilharz, 1852.
Syn.: Distoma hæmatobium, Bilh.; Distoma
capense, Harley, 1864.
Fig. 171.—Schistosoma
The Male is whitish, 12 to 14 mm. in hæmatobium, Bil.: male carrying
length, but is already mature when the female in the canalis
gynæcophorus. 12/1. (After
4 mm. long. The anterior end is
Looss.)
0·6 mm. or a little over in length. The
suckers are near each other, the oral
sucker is infundibular, and the dorsal
lip is longer than the ventral one. The
ventral sucker is a little larger,
0·28 mm., and is pedunculated. A
little behind the ventral sucker the
body broadens to a width of 1 mm., Fig. 172.—Transverse section
through a pair of Schistosoma
decreasing, however, in thickness; the
hæmatobium in copulâ. In the
lateral edges in-roll ventrally, so that male the point of reunion of the
the posterior part of the body intestinal forks has been cut
across. (After Leuckart.)
appears almost cylindrical, 0·4 to
0·5 mm. in diameter; the posterior extremity is somewhat more
attenuated. The dorsal surface of the posterior part of the body is
covered with spinous papillæ. There are delicate spines on the
suckers, and larger ones invest the entire internal surface of the
gynæcophoric canal, as well as a longitudinal zone at the edge of
that side of the external surface that is covered by the other side
rolling over it. The œsophagus is beset with numerous glandular
cells (fig. 173), and presents two dilatations; the intestinal
bifurcation is close in front of the ventral sucker, the two branches
uniting sooner or later behind the testes into a median trunk, which
may again divide at short intervals. The excretory pore is at the
posterior end, but placed somewhat dorsally; the genital pore is at
the beginning of the gynæcophoric canal, thus behind the ventral
sucker; into it opens the vas deferens which, posteriorly, broadens
into the seminal vesicle and then continues as the vasa efferentia of
the four or five testes (fig. 173).
The Female—filiform, about 20 mm. in
length, pointed at each end, and
measuring 0·25 mm. in diameter in the
middle. Their colour varies according to
the condition of the contents of the
intestine. (Posteriorly they are dark
brown or blackish.) The cuticle is
smooth except in the sucker, where
there are very delicate spines, and at
the posterior end, where there are
other larger spines. The oral sucker is a
little larger than the pedunculated
ventral sucker (0·07 and 0·059 mm.
Fig. 173.—Anterior end of the respectively). The anterior part of the
male Schistosomabody, 0·2 to 0·3 mm. in length; the
hæmatobium, Bilh. V.s.,
ventral sucker; I., gut cæca;
œsophagus is as in the male. The
G.p., genital pore; T., testes;intestinal bifurcation is in front of the
O.s., oral sucker; ventral sucker, the two branches
Oe.,
œsophagus with glandular uniting behind the ovary and the trunk
cells; V.s., vesicula seminalis.
running in a zigzag manner to the
40/1. (After Looss.)
posterior border. There are indications
of diverticula at the flexures. The ovary
is median. In young females it is of an elongated oval shape; in
older females the posterior end becomes club-shaped, whereas the
anterior end becomes attenuated; the oviduct originates at the
posterior end, but immediately turns forwards and joins the parallel
vitelline duct in front of the ovary (fig. 174), where the shell gland
cells open; the common canal becomes dilated to form the oötype,
and then proceeds as the uterus, with only slight convolutions, along
the central field to the genital pore, which lies in the middle line
immediately behind the ventral sucker. The single vitellarium starts
behind the ovary and extends to the posterior end. The acini are
situated at the sides of the excretory duct, which runs a median
course. The eggs are compact spindles, much dilated in the middle;
they have no lid, and are provided with a terminal spine
(rudimentary filament) at the posterior end, measuring 120 µ to
150 µ in length and 40 µ to 60 µ in breadth, but vary in size and
shape (fig. 175).
Distribution.—In order to understand the distribution of the worms
and eggs in the body, it may be well to recall the blood supply of the
abdominal and pelvic organs. It is generally assumed that the early
life (? cercarial stage) of the worms occurs in the liver, and that the
young worms travel from here, where they are invariably found, to
their various sites along the portal vein and its tributaries and so
against the blood stream. The tributaries of the portal vein are:—
(1) Superior mesenteric, the tributaries of
which are: (a) the veins of the small
intestine; (b) ileo-colic; (c) right colic; (d)
middle colic; (e) right gastro-epiploic; and
(f) inferior pancreatic. By these paths
infection of the small intestine, ascending
and transverse colon and pancreas would
occur.
(2) Splenic. (Ova have been recorded by
Symmers in the spleen.)
(3) Inferior mesenteric, the tributaries of
which are (a) superior hæmorrhoidal veins
from the upper part of the hæmorrhoidal
plexus; (b) sigmoid veins from sigmoid
flexure and lower portion of descending
colon; (c) left colic vein draining descending
colon.
The superior hæmorrhoidal veins form a
rich plexus in the rectum, and below this Fig. 174.—Schistosoma
hæma­tobium, Bilh.:
level in the upper and middle parts of the
genitalia of the female.
anal canal. The plexus forms two networks, V.s., ventral sucker; I.,
an internal plexus in the submucosa and an gut cæca; V.d., vitelline
external on the outer surface. The internal duct; V.sc., vitellarium;
plexus opens at the anal orifice into: (a) O., ovary; Oe.,
œsophagus; Sh., shell
branches of the inferior hæmorrhoidal vein
gland; U., uterus.
(from the pudic); (b) the external plexus. Magnified. (After
The external plexus gives off: (a) inferior Leuckart.)
hæmorrhoidal opening into internal pudic
(of internal iliac vein); (b) mid-hæmorrhoidal into internal iliac or its
branches; and (c) superior hæmorrhoidal opening into inferior
mesenteric. The external plexus further communicates with the
vesico-prostatic plexus. The vesico-prostatic (vaginal) plexus opens
into the vesical veins, which drain into the interior iliac vein. This
plexus also receives afferents from the pudendal plexus, the chief
tributary of which is the dorsal vein of the penis. The pudendal
plexus also receives branches from the inferior pudic and the
anterior surface of the bladder.
There is thus a communication between the portal vein and the vena
cava by means of these plexuses, viz., through the inferior and
middle hæmorrhoidals, and by the inferior hæmorrhoidals to the
bladder and thence by the vesical veins or the pudic to the caval
system (interior iliac).
It is thus by the inferior mesenteric and its tributaries that the
worms reach the descending colon, rectum, anal canal, and
eventually the bladder, and in some cases the caval system.
Before considering what is actually found post mortem in these veins
and the organs drained by them, we may further recall the fact that
the calibre of “medium” veins is 4 to 8 mm., “small” veins less than
40 µ in diameter and capillaries 8 µ to 20 µ. Further, the maximum
diameter of the male worm is 1 mm., that of the female 280 µ and
eggs in utero 80 µ to 90 µ long by 30 µ to 40 µ.
Liver and Portal Vein.—Here worms are most easily found post
mortem. Often only males are found and these of the same size, and
if females occur only a few worms are found in copulâ. The worms
are frequently not full size and the males may contain no free
spermatozoa in their testes, and as regards the females some may
be fertilized, others not, as shown by the presence or absence of
spermatozoa in the seminal receptacle or uterus. In either case they
may contain eggs—lateral-spined—usually one, less often two, but
there may be as many as five or six. These eggs may also show
some abnormality, which takes the form of: (1) abnormal contents,
viz., disintegrating yolk cells with or without an ovarian cell; (2)
abnormal shape but with normal contents and probably represented
by the collapsed and empty egg-shells which are found in the
tissues.
As to the interpretation of these facts, Looss believes that these
lateral-spined eggs are products of young females whose egg-laying
is not at first properly regulated. The shape that the eggs take, viz.,
with a lateral spine, is determined by an excess of material—ovarian
and yolk cells—being present in the oötype. The shape of eggs
depends upon the position they have in the oötype during their
formation. In young females an excess of cells—yolk cells especially
—accumulates, distending not only the dorsal wall but a portion also
of the short duct joining the oötype to the uterus. The result of this
is that the axis of the oötype and egg is almost transverse to the
body, and the posterior funnel-shaped portion of the oötype, instead
of being terminal, has now a lateral or rather a ventral position, so
that the spine which occupies this portion, instead of being terminal,
is now lateral. It is noteworthy that these lateral-spined eggs are
thicker, owing to the excess of material present, and not
uncommonly have a curved anterior border, due to a projection of
the anterior end into the anterior opening of the oötype.
As these eggs are being laid by females in the portal vein they are
carried back to the liver by the blood stream. The liver is one of the
commonest sites for these eggs; also terminal-spined eggs may be
found here for the same reason.
Hæmorrhoidal Veins.—Mature worms, generally in copulâ, are
usually found here, though young not fully grown females may also
occur. The tissues of the rectal wall (or colon) show, as a rule, large
quantities of lateral-spined eggs, though less often only terminal-
spined eggs may be found.
Vesico-prostatic Plexus.—Worms in copulâ are found in the veins of
the submucosa in the bladder, and the eggs in the mucosa, and
those voided are usually terminal-spined, though lateral-spined eggs
are not so rare as generally thought. The problem next arises as to
how the eggs get to the lumen of the gut or bladder.
The female worm is 280 µ in diameter. Veins in the submucosa of
the rectum less than 178 µ in diameter are not affected with
endophlebitis. It is probable that the female even by stretching could
not penetrate much beyond this. Eggs are probably then laid in the
submucosa as near the muscularis mucosa as possible. Now if the
eggs are laid in a vein of larger calibre than the worm fills, the eggs
would be carried back to the inferior mesenteric vein, so that
presumably the worm must succeed in blocking the vein already
narrowed by endophlebitis, so that by the stasis which ensues the
eggs may escape from the veins. How this occurs is not exactly
known; it is not necessarily due to the spine, as the same escape
into the tissues occurs in spineless eggs, such as those of
Schistosoma japonicum. The eggs, then, pass as foreign bodies
through the tissues. Another hypothesis is that the worms leave the
veins in order to lay their eggs, but the evidence is against this.
Caval System.—Occasionally worms that have passed through the
vesical plexus may be found in the iliac vein, inferior vena cava, and
even the lungs. If the worms are young they contain a lateral-spined
egg; if adult, numerous (50 to 100) terminal-spined eggs.
Lungs.—When the liver is strongly infected with (terminal-spined)
eggs it is possible that by passive movements some may pass into
the intralobular veins, and thence by the inferior vena cava to the
lungs.
Gall-bladder.—Similarly terminal-spined eggs pass into the bile-
capillaries and gall-bladder (where they may be abundant), and so
into the fæces.
Detection of Eggs.—Occasionally eggs may be found in various other
parts of the body. They are best detected by macerating pieces of
 1
the tissue in question in about 4
per cent. hydrochloric acid at 50 to
60°C. (Looss).
Pathological changes:—
Rectum.—These have been studied thoroughly by Letulle in the case
of an apparently pure infection of the rectum. 274 They take the form
of a chronic diffuse inflammation, which may result in—(1)
ulceration, or (2) hyperplasia of the mucosa, producing adenomata.
Ulcerative Form.—The mucosa is transformed into a mass of vascular
connective tissue. The connective tissue spaces next become
invaded by numerous mononuclear cells. The tissue itself undergoes
diffuse sclerosis, becoming hard and fibroid. Eventually ulcerative
necrosis sets in. During these changes the Lieberkühn glands are
destroyed. The process does not extend to the submucosa, in this
respect differing from that in chronic dysentery.
Hyperplastic Form.—The Lieberkühn glands of the mucosa at first
hypertrophy; then there is an actual hyperplasia resulting in
adenomata. The interstitial tissue of the glands is also greatly
hypertrophied, giving rise to very vascular granulations. These
growths are often hollow and contain worms. Many eggs are found
in the mucosa on their way to the lumen of the gut.
The muscularis mucosa is thickened up to twice or even ten times
the normal. Its vessels are dilated (36 µ to 80 µ), but they do not
allow of the passage of worms.
The submucosa is profoundly changed; rigid and hard instead of
supple. It is here that the greatest number of eggs occur. A
remarkable condition of endophlebitis exists in the veins of the
submucosa, not only in the smaller ones but also in the larger ones
(370 µ by 270 µ). This endophlebitis results in a more or less
complete occlusion of the vessels of the lumen.
The muscular coats are free from change, also their veins.
The Serous Coats.—The veins about 1,900 µ, also show
endophlebitis. Besides the rectum, in extreme cases even the
transverse colon, the cæcum and small intestine may be affected.
Bladder.—In the early stages the mucosa is deep red and swollen
like velvet, or there may be localized patches of hyperæmia or
extravasation. The subsequent changes take two chief forms:—
(1) Sandy Patches.—The mucosa looks as if it were impregnated
with a fine brownish or yellowish powder (myriads of ova). This is
accompanied by a gradual hypertrophy and new formation of
connective tissue, so that dry, hard or plate-like patches with this
sandy appearance arise; the thickening eventually affects all the
coats of the bladder. In the older patches many of the eggs are
calcified. These patches sooner or later break down, ulcerate and
necrose. Phosphatic deposits are abundant and stone is common.
These patches are not found in the rectum.
(2) Papillomata.—Where the inflammatory change produced by the
eggs gives rise to hypertrophy and hyperplasia of the mucosa,
papillomata result, the axis of which is formed by connective tissue
of the submucosa. These are most variable in shape and form and
bleed readily, and sometimes contain cavities of extravasated blood.
As in the rectum, it is in the submucosa that eggs are most
abundant, and worms in copulâ occur in the veins of this layer, but
endophlebitis is not as general as described in the rectum. Malignant
disease of the bladder is not an uncommon sequela of bilharziasis.
Besides the bladder, the ureters and kidneys may in advanced cases
be involved. The prostate and vesiculæ seminales are commonly
diseased. Eggs have been recorded in the semen. The urethra is
frequently attacked; the vagina in the female.
Eggs also occur in the lymphatic glands of the gut.
Geographical Distribution.—East Africa: Nile Valley, Red Sea Coast,
Zanzibar, Portuguese East Africa, Delagoa Bay, Natal, Port Elizabeth.
South Africa: Cape Colony, Orange Free State, Transvaal, Mauritius,
Bourbon, Madagascar.
West Africa: Angola, Cameroons, Gold Coast, Gambia, Senegal,
Sierra Leone, Lagos, Nigeria.
North Africa: Tripoli, Tunis, Algeria, parts of the Sahara.
Central Africa: Sudan, various portions. Uganda, Nyasaland.
It occurs with varying frequency in these regions. It is probably more
widely spread than this list implies, as undoubtedly many cases are
seen which are not recorded.
Isolated cases have been recorded from Arabia, India, 275 Greece,
Cyprus.
The means by which infection is brought
about are still uncertain; we only know that
the miracidia (fig. 175) enclosed in the
discharged eggs do not hatch if the eggs
remain in the urine, but after cooling perish.
As soon, however, as the urine is diluted with
water the shell swells, generally bursting
lengthways, and releases the miracidium from
its investing membrane, so that it can swim
about with the aid of its cilia. In its structure it
differs but little from the miracidium of
Fasciola hepatica, as, for instance, in the lack Fig. 175.—Ovum of
of eyes; the two large gland cells situated on Schisto­soma hæma­to­‐
either side of the intestinal sac are also bium, Bilh., with mira­‐
cidium, which has
present in the miracidia of Fasciola hepatica.
turned its anterior end
Sarcode Globules.—This is a term applied to towards the posterior
certain globules which at times appear in the
miracidium and are later ejected. Some end of the egg. 275/1.
authors consider them as indicative that the (After Looss.)

miracidium has developed into a sporocyst,

but Looss considers them to be degeneration products.
The Bilharzia mission, under R. T. Leiper, sent to Egypt by the War
Office early in 1915, reports that cercariæ of bilharzia type were
recognized in four of the commonest fresh-water molluscs around
Cairo.
With material obtained from naturally infected Planorbis boissyi
acute bilharziosis was experimentally produced in rats, mice, and
monkeys. Infection takes place experimentally through the skin and
also through the mucous membrane of the mouth and œsophagus.
The miracidium, after entering the mollusc, develops into a
sporocyst. This gives rise not to rediæ, but to secondary sporocysts,
which, in turn, produce cercariæ. These, like the adult worm, differ
from other distomes in lacking a muscular pharynx.

Schistosoma mansoni, Sambon, 1907.

According to Manson, Sambon and others, the eggs with lateral
spines belong to a species different from Schistosoma hæmatobium.
Infections with this species only are said to occur in the Congo,
Southern States of North America, West Indies (Guadeloupe) and
Brazil (Bahia). The following characters, according to Flu,
differentiate this species: (1) In the male the transition from the
anterior portion of the worm to the lateral fields (the infolded
portions which form the gynæcophoric canal) is not a gradual one as
in Schistosoma hæmatobium, but in this case the lateral fields rise
suddenly, almost at right angles to the anterior portion. (2) The
ovaries have a well-marked convoluted course as in no other
schistosome. (3) The oötype is symmetrical in reference to the long
axis of the body, its duct being lateral on the ventral side (Looss’
explanation of this we have already given). (4) The worms live
exclusively in portal vein and tract. (As lateral-spined eggs occur also
in the bladder, this is not exactly true.)

Schistosoma hæmatobium, Bilharz, 1852.

Male, four or five large testes. Gut forks unite
late, so that the single gut stem is short.
Female, ovary in posterior half of body. Uterus
very long, voluminous, with many terminal-
spined eggs, some lying in pairs. Vitellaria in
posterior fourth of body. Cercariæ in Bullinus
contortus and Bullinus dybowski (syn.: Physa
alexandrina) in Egypt.

Schistosoma mansoni, Sambon, 1907.

Male, eight small testes. Gut forks unite early,
so that the single gut stem is very long.
Females, ovary in anterior half of body. Uterus
very short; usually only one lateral-spined egg
at a time in utero. Vitellaria occupy posterior
two-thirds of body. Cercariæ in Planorbis
boissyi in Egypt.
The above morphological descriptions are
founded on worms of each species, derived
from experimentally infected mice (Leiper,
R. T., Brit. Med. Journ., March 18, 1916,
p. 411).

Schistosoma japonicum, Katsurada, 1904.

 Syn.: S. cattoi, Blanchard, 1905.

Male.—Eight to 19 mm., but extreme limits are 5

to 22·5 mm. Consists of a short fore-body,
separated by the ventral sucker from the hind-
body. The ventral sucker is stalked and
somewhat larger than the oral sucker. Both
suckers are larger than the corresponding ones
in S. hæmatobium. Body usually smooth, but in
the fresh state numerous fairly evident spines
along the margin of the canal. Œsophagus: two
bulbs. The junction of the gut forks more
posterior than in S. hæmatobium, the median
united gut stem occupying a quarter to one-fifth
to one-sixth of the body length. An excretory
canal runs along each side of the body, opening
into the dorsal excretory pore. Testes irregularly
elliptical, six to eight in number, in the anterior
part of hind-body. The vasa efferentia unite into
a common vas deferens which opens directly
behind the ventral sucker. The seminal vesicle
lies just behind this.
Female.—Up to
26 mm., generally
thinner than the male.
Surface smooth.
Suckers armed with Fig. 176.—
fine spines. Ventral Schistosoma
Fig. 177.—Schistosoma
japonicum: anterior
japo­ni­cum, male and sucker larger than end with testes;
female in copulâ. × 60. oral. Body thicker posterior end with
(After Katsurada.)
behind the region of point of union of
the ovary. The gut cæca. Length of
forks unite immediately behind the ovary. The worm about
united gut much thicker than in S. 10 mm. (After
hæmatobium. Ovary elliptical, almost in the Katsurada.)
mid-body, its hinder portion dilated. The oviduct
arises from its posterior end and then runs sinuously forward, where
it is joined by the vitellarian duct; the vitellarium well developed,
extending from behind the ovary almost but not quite to the
posterior end as in S. hæmatobium. Shell gland ducts enter at the
junction point of oviduct and vitelline duct. The canal here forms an
oötype and then proceeds as the uterus to open directly behind the
ventral sucker. The uterus occupies almost half the hind-body. In S.
hæmatobium this is not so. The uterine canal is cleft-like, i.e., its
dorso-ventral diameter is much greater than its lateral diameter. The
number of eggs varies from about 50 to 300 from observations
made in various hosts.
Eggs.—In utero assume various shapes, as they are soft; the lumen
of the uterus is narrow. Outside they are oval, faint yellow, double
contoured. In fæces the eggs measure 83·5 µ, by 62·5 µ (man);
85 µ by 61·5 µ (cattle); 98·2 µ by 73·8 µ (dog). The eggs have
either small lateral spines or thickenings, and Looss at the opposite
side has described cap-like thickenings. The eggs in the tissues
undergo various deformities, and may contain a miracidium, as also
the eggs in fæces do; or the contents may consist of granular matter
or amorphous masses or they may be calcified. Lymphocytes and
giant cells may also invade the eggs.
 Fig. 178.—Schistosoma japonicum: eggs from human liver,
showing “spines” and “hoods” at opposite pole. (After Looss.)

Mode of Infection.—The miracidia hatch in

water in as little as fifteen minutes, but the
majority in one to three hours. They will live
in water for about twenty-four hours. In
water they undergo a transformation into
“larvæ,” which then penetrate the skin, as
has been shown by Japanese writers to
hold good for man, cattle, dog and cat. The
penetration of the skin is attended with an
eruption on the legs, “Kabure.” The exact
route by which the worms reach the portal Fig. 179.—Schistosoma
vein is uncertain. Infection in Japan takes japo­ni­cum: from dog.
place from spring to autumn, especially May Uterine egg. × c. 800.
(After Katsurada.)
to July, when the soil is contaminated with
manure of cattle infected with S.
japonicum. They also appear to develop in molluscs. Leiper and
Atkinson found cercariæ (in sporocysts) in the liver of a mollusc,
Katayama nosophora. They infected mice by immersing them in
water containing liver emulsion and so free cercariæ, thus
confirming the similar results of Miyairi and Suzuki.
Habitat.—The worm occurs in Japan, China, and the Philippines. The
normal host is man and mammals. Cattle, dog and cat are often
found naturally infected. Mice can also be experimentally infected.
Their seat of election is the portal vein and its branches, especially
the mesenteric veins. They either swim free in the blood or remain
fixed by their suckers to the intima of the vessels. They have also
been found in the vena cava and right heart of a cat, but not so far
in the vesical plexus.

Fig. 180.—Schistosoma Fig. 181.—Schistosoma

japonicum: from dog. × c. japonicum: from dog. Egg
800. (After Katsurada.) from fæces. × c. 800. (After
Katsurada.)

Eggs are found in the submucosa and mucosa of the gut, especially
the colon, and at times in the serosa and subserosa of the small
intestine, where they give rise to new growths. Occasionally eggs
are found in the brain. The life of the worms is at least two years.
Pathogenic Effects.—Anæmia through loss of blood due to worms;
enlarged spleen, toxic in origin (?); phlebitis, thrombosis, due to
portal stasis; the eggs, however, cause the greatest mischief. They
are carried by the circulation to various organs where they produce
inflammation, granulation tissue, and later connective tissue.
Liver.—The eggs reaching this organ give rise to granulomata and
hence enlarged liver, and later, when connective tissue is formed, to
contraction. The surface is rough and irregularly granular, “parasitic
embolic cirrhosis” of Yamagiwa.
 Fig. 182.—Schistosoma japonicum: section through the gut of a
Chinaman showing eggs. × 58. (After Catto.)

Gut.—The eggs in the mucosa and submucosa cause catarrh and

destruction of tissue or new growth. In the small intestine the eggs
are mainly in the serosa and subserosa, where they give rise to
polypoid or branched growths.
Spleen.—Enlarged, at first due to toxin (?) and later due to portal
stasis. Eggs in the spleen are uncommon.
Ascites also arises from the portal stasis, and is generally present in
advanced cases.
Eggs may be found in many other situations: glands (numerous),
mesentery, stomach, pancreas, kidney, etc. The bladder remains
free.

Fig. 183.—Schistosoma japonicum: liver showing eggs in the intra- and

interlobular connective tissue. × c. 80. (After Katsurada.)

Class III. CESTODA, Rud., 1808.

Tapeworms have been known from ancient times—at all events, the large species
inhabiting the intestines of man—and there has never been a doubt as to their
animal nature. The large cysticerci of the domestic animals (occasionally of man
also) have been known for an equally long period, but they were generally
regarded as growths, or “hydatids,” until almost simultaneously Redi in Italy, and
Hartmann and Wepfer in Germany, concluded from their movements and
organization that they were of animal nature. From that time the cysticerci have
been included amongst the other intestinal worms, and Zeder (1800) established a
special class (Cystici, Rud., 1808) for the bladder worms. Things remained in this
condition until the middle of the last century, when Küchenmeister, by means of
successful feeding experiments, demonstrated that the cysticerci were definite
stages of development of certain tapeworms. Before Küchenmeister, E. Blanchard,
van Beneden, and v. Siebold had held the same opinion in regard to other asexual
Cestodes.
Since the most remote period another question has again and again occupied the
attention of naturalists, the question of the morphological nature—that of the
individuality of the tapeworm. The ancients, who were well acquainted with the
proglottids (Vermes cucurbitani) that are frequently evacuated, were of the
opinion that the tapeworm originated through the union of these separate
proglottids, and this view was maintained until the end of the seventeenth century.
In 1683 Tyson discovered the head with the double circlet of hooks in a large
tapeworm of the dog; Redi (1684) was also acquainted with the head and the
suckers of several Tæniæ. Andry (1700) found the head of Tænia saginata, and
Bonnet (1777) and Gleichen-Rusworm (1779) found the head of Dibothriocephalus
latus. Consequently most authors, on the ground of this discovery, considered the
tapeworm as a single animal, that maintains its hold in the intestine by means of
the head, and likewise feeds itself through it. The fact was recognized that there
were longitudinal canals running through the entire length of the worm, and it was
thought that these originated in the suckers, and that the entire apparatus was an
intestine. As, moreover, the segments form at the neck, and are cast off from the
opposite extremity, the tapeworm was also compared with the polyps, which were
formerly regarded as independent beings.
Steenstrup, in his celebrated work on the alternation of generations (1841), was
the first to give another explanation. This has been elaborated still further by van
Beneden, v. Siebold and Leuckart, and until a few years ago all authorities adopted
his views. According to this view, the tapeworm is composed of numerous
individuals, something like a polyp colony, and, in addition to the proglottids—the
sexual individuals which are usually present in large numbers—there is one
individual of different structure, the scolex, which not only fastens the entire
colony to the intestine, but actually produces this colony from itself, and therefore
is present earlier than the proglottids. The scolex is a “nurse,” which, though itself
produced by sexual means, increases asexually like a Scyphistoma polyp; the
tapeworm chain has therefore been termed a strobila. Consequently the
development of the tapeworms was explained by an alternation of generations. In
support of this opinion it was demonstrated not only that the adult sexual
creatures, the proglottids, can separate from the colony and live independently for
a time, but that in certain Tæniæ, and especially in many Cestodes of the shark,
the proglottids detach themselves long before they have attained their ultimate
size, and thus separated continue to develop, grow and finally multiply; the scolex
also exhibits a certain independence in so far as, though not, as a rule, capable of
a free life, yet it in some cases lives as a free being, partly on the surface of the
body of marine fishes and partly in the sea. With the more intimate knowledge of
the development of the cysticerci, the independent nature of the scolex was
recognized. It is formed by a budding of the bladder that has developed from the
oncosphere, in some cases (Cœnurus) in large numbers, in other cases
(Echinococcus) only after the parent cyst has developed several daughter cysts.
Released from its mother cyst and placed in suitable conditions, it goes on living,
and gives rise at its posterior end by budding to the strobila, the proglottids of
which eventually become sexual individuals.
In order to make this clearer we will briefly summarize what takes place in the
jelly-fishes.
By metamorphosis is meant a developmental change in the same individual, while
alternation of generations, or metagenesis, implies a stage in which reproduction
of individuals takes place by a process of budding or fission. This asexual
reproductive stage alternates with the sexual mode of reproduction. Thus in the
development of the Scyphozoa (jelly-fishes) we have:—
(1) The fertilized egg cell divides regularly and forms a morula.
(2) By accumulation of fluid in the interior this becomes a closed sac with a wall
formed of a single layer of cells, forming the blastosphere or blastula.
(3) One end of the sac is invaginated, forming a gastrula.
(4) The gastrula pore or mouth closes, forming again a sac, the walls of which
have two layers, forming a planula.
(5) This becomes fixed to a rock, an invagination forms at one end, a depression—
the stomodæum—communicating with the enteric cavity. Tentacles grow out and
we have a Scyphozoön polype, Scyphistoma or Scyphula. It is to this stage that
Steenstrup gave the name “nurse” (“wet-nurse”), because it nourished or
produced asexually the succeeding forms.
(6) Asexual reproduction by transverse fission occurs in this, forming a pile of
saucer- or pine-cone-like animals which before this time had been considered to
be a distinct animal, which was called strobila from its resemblance to a pine-cone.
This is the alternate generation.
(7) The individuals of the strobila become free and are called Ephyrulæ.
(8) These develop finally into adult sexual jelly-fish, Scyphozoa, so that comparing
a tapeworm with this we have (a) egg, (b) scolex (= Scyphula or “nurse”), (c)
asexual reproduction of the tapeworm chain (= strobila), (d) development of the
individuals of the chain (proglottids) into sexual adults.
Van Beneden’s terminology for these stages is the following: Ciliated embryo =
protoscolex; scyphistoma = deutoscolex (or scolex); free Ephyrula = proglottis.
According to this view, as is the case in many endoparasitic Trematodes, asexual
reproduction by budding occurs at two stages of the whole cycle of development,
viz. (1) in the formation of the scolex by budding from the bladder (“nurse”), (2)
in the formation of the strobila by budding from the scolex (“nurse”).
But in cysticercal larval forms it appears that the scolex does not arise in this way
but is simply a part of the proscolex (hexacanth embryo), becoming invaginated
into it for protection, so that there is no asexual gemmation here. It has been
questioned also whether the strobila also arises by gemmation. If it does, the
tapeworm is a colony of zoöids produced by budding from the asexual scolex; if it
is not produced in this way, then the tapeworm is to be regarded as an individual
in which growth is accompanied by segmentation. Against the “colony” view are
the facts that the muscular, nervous, and excretory systems are continuous
throughout the worm, and that some tapeworms, such as Ligula, are
unsegmented.
Finally, if the tapeworm is an individual the question arises which is the head end.
As new segments are formed at the neck, and as this point in annelids is the
antepenultimate segment, the scolex must be the last or posterior segment. The
caudal vesicle or bladder of larval forms is consequently anterior. According to this
view, in tapeworms as among many endoparasitic flukes, an asexual multiplication
occurs at two points of the whole cycle of development, which is as follows: (1)
egg, (2) oncosphere or hexacanth embryo, (3) bladder (cysticercus or hydatid),
(4) (after digestion of the bladder) by budding, the scolex, (5) by budding from
the scolex the sexual proglottids, (6) the egg; (4) and (5) being the two asexual
stages.

Anatomy of the Cestoda.

If we except the tapeworms with only one proglottis, the Cestoidea

Monozoa, Lang = Cestodaria, Monticelli, we can always distinguish in
the Cestodes, in the narrower sense, one scolex or head and a large
or small number of segments (proglottids). The scolex serves the
entire tapeworm for fastening it to the internal surface of the
intestinal wall, and therefore carries at its end various organs which
assist in this function, and which are as follows: (i) Suctorial organs,
i.e., the four suckers (acetabula), which are placed crosswise at the
circumference of the thickened end of the scolex; further, the double
or quadruple groove-like suckers (bothridia), which are diversely
shaped in the various genera and families. 276 (2) Fixation organs
(hooklets) 277 that likewise occur in varying numbers and different
positions; they may be in the suckers, or outside them on the apex
of the scolex; for instance, in many of the Tæniidæ they appear in a
circle around a single protractile organ, the rostellum, or the latter
may be rudimentary, and is then replaced by a terminal sucker. (3)
Proboscis. One family of the Cestodes, the Rhynchobothriidæ, carries
four proboscides, moved by their own muscular apparatus, on the
scolex, and they are beset with the most diverse hooks. (4) Tentacle-
like formations are only known in one genus (Polypocephalus).

The thickened part of the scolex that carries the suckers is usually
called the head; the following flat (unsegmented) part connecting it
with the proglottids is called the neck, and is sometimes quite small.
In a few cases the entire scolex (or head) disappears, and its
function is then undertaken by the contiguous portion of the chain of
proglottids, which is transformed into a variously shaped pseudo-
scolex.

The proglottids are joined to the scolex in a longitudinal row, and are
arranged according to age in such a manner that the oldest
proglottis is farthest from the scolex, and the youngest nearest to it.
The number of segments varies, according to the species, from only
a few to several thousands; they are either quadrangular or
rectangular; in the latter case their longitudinal axis falls either
longitudinal or transverse to that of the entire chain, according as
the segments are longer than broad or broader than long. When the
number of segments is very large, the youngest ones are, as a rule,
transversely oblong, the middle ones are squarish, and the mature
ones longitudinally oblong. The posterior border of the segments, as
a rule, carries a longitudinal groove for the reception of the shorter
anterior border of the following proglottis. The two lateral borders of
the segment are rectilinear, but converge more or less towards the
front, or they are bent outwards. In most of the Cestodes the
segments, just as the neck, are very flat; in rare cases their
transverse diameter is equal to their dorso-ventral diameter. As a
rule the segments, singly or several united together, detach
themselves from the posterior end, in many cases only after
complete maturity is attained, and in others much earlier; they then
continue to live near their parent colony, to still call it by that name,
in the same intestine and continue their development. Even when
evacuated from the intestine the proglottids under favourable
circumstances can continue to live and creep about, until sooner or
later they perish.
The first proglottis formed, and which in a complete tapeworm [i.e.,
sexually complete] is the most posterior, is as a rule smaller and of
different shape, it also frequently remains sterile, as likewise
happens in the next (younger) segments in a few species;
otherwise, however, sooner or later the generative organs develop in
all the segments, mostly singly, sometimes in pairs; in the latter case
they may be quite distinct from each other or possess some parts in
common. The term “mature” is used for a proglottid that has the
sexual organs fully developed, while “gravid” is used for one
containing eggs. Most of the species combine male and female
genitalia in the same segment, only a few are sexually distinct
(Diœcocestus). In the hermaphrodite species one male and one
female sexual orifice are always present, and, in addition, there may
be a second female orifice, the uterine opening; as a rule, however,
this is lacking, and in one sub-family, the Acoleinæ, to which also the
genus Diœcocestus belongs, the other sexual orifice, the opening of
the vagina, is also absent. The position of these orifices varies; the
cirrus and vagina usually open into a common atrium on one lateral
border or on a surface of the segments; the orifice of the uterus may
be on the same surface or on the opposite one.
The surface on which the uterus opens is termed the ventral surface;
if this orifice is absent, one must depend on the ovary, which almost
always approaches one of the two surfaces; this surface is then
called the ventral.
The length of the Cestodes—independently of their age—depends on
the number and size of the segments, as well as on their
contraction; the smallest species (Davainea proglottina) is 0·5 to
1·0 mm. in length; the largest may attain a length of 10 m., and
even more.
The entire superficial surface of the tapeworms is covered with a
fairly resistant and elastic layer, which exhibits several indistinctly
limited layers and which is usually called a cuticle, which also covers
the suckers, and is reflected inwardly at the sexual orifices. In some
species fine hairs appear, either on the entire body or only in the
region of the neck, on the external surface. In the cuticle there can
be recognized, besides the pores, which no doubt are concerned
with nutrition, spaces in which lie the ends of sensory cells. Close
under the cuticle lies the external layer of the parenchyma (basal
membrane), and below this the circular and longitudinal muscles
forming the dermo-muscular coat. The matrix cells of the cuticle
occur as in the Trematodes, only on the inner side of the peripheral
muscles in the external zone of the parenchyma; they are fusiform
cells, forming one or two layers, but are not arranged in the manner
of epithelial cells (fig. 184, Sc.c.). They have fine branching
processes which run between the dermal muscles, pass through the
basal membrane and penetrate the internal surface of the cuticle
with small pistil-like enlargements, expanding on the internal surface
of the cuticle into a thin plasma layer.
In addition to the above mentioned,
there are other cuticular formations
occurring on the cuticle of some
Cestodes, such as immobile hairs and
variously formed hooks, such as are
seen principally on the scolex. Their
development is only roughly known in
a few species; they are usually
already present in the larval stage,
and of the same arrangement and
shape as in the fully developed
tapeworms; a matter of importance,
because by these structures larvæ
can be recognized as being those of a
certain species of tapeworm.
The cuticular glands in Cestodes are
scarce.
Fig. 184.—Schematic
The parenchyma forms the chief tissue representation of a small part of
of the entire body, and in all a transverse section of Ligula sp.
essentials its structure is similar to Bs., basal membrane; Cu.,
cuticle; at its base are the
that of the Trematodes.
endplates of the subcuticular
The same doubt exists here also as to (epithelial) cells; in the centre a
the nature of the parenchyma. Recent cuticular sense organ, O.s.;
F.v.s., vitelline follicle; Exc.,
authors consider that it consists of excretory vessel; C., calcareous
highly branched cells, the processes corpuscle; L.m., longitudinal
of which ramify in all directions. muscles; M.c., myoblast; P.m.,
These cells lie in a non-cellular matrix parenchymatous or dorso-ventral
containing fluid vacuoles. This matrix muscles; Pl., plexus of nerve
fibres; A.m., circular muscles;
Sc.c., subcuticular or matrix cell;
spreads in between and so breaks the T.c., terminal flame cell. 500/1.
continuity of the epidermal cells. (After Blochmann.)

In the parenchyma of almost all the Cestodes there are found in

adult specimens, as well as in larvæ, light-refracting concentrically
striated structures, of a spherical or broad elliptical shape, which, on
account of their containing carbonate of lime, are termed calcareous
corpuscles (fig. 184, C.). Their size, between 3 µ and 30 µ, varies
according to the species; their frequency and distribution in the
parenchyma also varies, but they are chiefly found in the cortical
layer. They are the product of certain parenchymatous cells, in the
interior of which they lie like a fat globule in a fat cell, but according
to others they are intercellular in origin.
The muscular system of the proglottids is composed of—(1) the
subcuticular muscles (figs. 184 and 185), as a rule consisting of a
single layer of annular muscles; (2) longitudinal muscles; (3) dorso-
ventral fibres extending singly from one surface to the other, and at
both ends expanding in a brush-like manner, and inserted into the
basal membrane, consisting of an outer, more numerous, and an
inner, less numerous but more powerful layer (the number of
bundles in this layer being in certain cases of specific importance);
(4) transverse fibres, the elements of which penetrate to the borders
of the segments, thus passing through the longitudinal muscles and
reaching the cuticle. In the region of the septa the transverse and
dorso-ventral muscles form a kind of plate.
 Fig. 185.—Half of a transverse section through a proglottis of Tænia
crassicollis. Cu., cuticle; Ex.v., external excretory vessel, to the right of
which there is the smaller internal one; T., testicular vesicles; L.m.,
longitudinal muscles (outer and inner); M.f., lateral nerve with the two
accessory nerves; Sc.c., subcuticular matrix cells; Sm.f., submedian nerve;
Tr.m., transverse muscles; Ut., the uterus, and the middle of the entire
transverse section. 44/1.

The mass of parenchyma bounded by the transverse muscles is

termed the medullary layer, while the mass lying outside them is
termed the cortical layer.
It was known long ago that the myoblasts adhere to the dorso-
ventral fibres as thickenings, but it is only recently that large star-
shaped cells (fig. 184), separated from but connected with them by
processes, have been recognized as the myoblasts of other fibres
(Blochmann, Zernecke).
Within the scolex the direction and course of the muscular layers
change.
The suckers are parts of the musculature, locally transformed, with a
powerful development of the dorso-ventral muscles, now become
radial fibres.
The rostellum of the armed Tæniæ, like the proboscis of the
Rhynchobothriidæ, also belongs to the same category of organs.
In the simplest form, the rostellum,
or top of the head (as in Dipylidium
caninum), appears as a hollow oval
sac, the anterior part of which,
projecting beyond the upper surface
of the head, carries several rows of
hooks (fig. 186). The entire internal
space of the sac is occupied by an
elastic, slightly fibrous mass, while
the anterior half of the surface of the
rostellum is covered by longitudinal
fibres and the posterior half by Fig. 186.—Dipylidium caninum:
from the cat. In the upper figure
circular fibres. On contraction of the the rostellum is retracted, in the
latter the entire mass is protruded lower protruded, a, sucker; b,
through the apical aperture, the hooks of rostellum; B, enlarged
surface of the rostellum becomes hook; c, apical aperture on
more arched, and the position of the scolex; d, longitudinal muscles;
e, circular muscles. (After
hooks is, in consequence, altered. Benham.)
The rostellum of the large-hooked
Tæniidæ, which inhabit the intestine
of man and beasts of prey, is of a far more complicated structure,
for, in addition to the somewhat lens-shaped rostellum carrying the
hooks on its outer surface, there are secondary muscles grouped in
a cup-like manner (fig. 187). Every change in the curvature of the
surface of the rostellum induces an alteration in the position of the
hooks. In the hookless Tæniidæ the muscular system of the
rostellum is altered in a very different manner; in a few forms a
typical sucker appears in its place.
The nervous system commences in the scolex and runs through the
neck and the entire series of proglottids. Within the proglottids it
consists of a number of longitudinal nerve fibres of which those at
each lateral border are usually the largest. In the Tæniæ the lateral
 nerves are accompanied both dorsally
and ventrally by a thinner nerve
(accessory nerve) (fig. 185); on each
surface, moreover, between the
lateral nerve and the median plane,
there are two somewhat stronger
bundles (sub-median), so that there
is a total of ten longitudinal nerve
bundles. They lie externally to the
transverse muscle plates, and the
lateral and accessory bundles lie
externally to the principal excretory
vessels, and are everywhere
connected by numerous anastomoses
Fig. 187.—Longitudinal section of
the head and neck of Tænia and secondary anastomoses; one
crassicollis, showing the lens- typical ring commissure is usually
shaped muscular rostellum, with found at the posterior border of the
two hooks lying in the concentric segments. In the Bothriocephalidæ
cup-like mass of muscles. L.m.,
longitudinal muscles of the neck;
the distribution of the nerve bundles
L.f., left lateral nerve; G., is different (for instance, two lie in
ganglion; S.c., subcuticular the medullary layer), or they are split
layer; W1, external, W2, internal up into a larger number of branches.
excretory vessel. 30/1. In the scolex the nerve bundles are
connected in a very remarkable
manner by commissures with that which is generally termed the
central part of the entire nervous system. There occurs normally a
commissure between the two lateral nerves; at the same level, the
dorsal and ventral median nerves are also connected at each surface
as well with each other as with the lateral nerves, so that a
hexagonal or octagonal figure is formed. The so-called apical nerves
pass from this commissural system anteriorly, embrace the
secondary muscular system of the rostellum semicircularly, and form
an annular commissure (rostellar ring) at the inner part of the
rostellum.
The peripheral nerves arise from the
nerve bundles as well as from the
commissures situated in the scolex;
some go direct to the muscles, while
others form a close plexus of nerves
external to the inner longitudinal
muscles, which plexus likewise sends
out fibres to the muscles, but
principally to numerous fusiform
sense organs (fig. 184, Pl.); they lie
internal to the subcuticular cells and,
piercing the cuticle with their
peripheral processes, end as
projecting “receptor” hairs. Higher
organs of sense are not known.
Fig. 188.—Tænia cœnurus, head
The excretory apparatus of the and part of neck showing
Cestodes is similar to that of other nervous system. Enlarged. (After
flat worms. The terminal (flame) Niemiec.)
cells, which hardly differ in
appearance from those of the Trematodes, are distributed
throughout the parenchyma, but are more common in the cortical
than in the medullary layer (fig. 184, T.c.). Before opening into a
collecting tube, the capillaries run straight, tortuously, or in
convolutions, anastomosing frequently with one another or forming
a rete mirabile. The collecting tubes, which have their own epithelial
and cuticular wall, and which also appear to be provided with
muscular fibres, occur typically as four canals passing through the
entire length of the worm (fig. 189); they lie side by side, two (a
wider thin-walled ventral, and a narrower thick-walled dorsal one) in
either lateral field; in the head the two vessels on each side unite by
means of a loop, at the posterior extremity they open into a short
pyriform or fusiform terminal bladder which discharges in the middle
of the posterior edge of the original terminal proglottis.
This primitive type (fig. 189) of arrangement of collective tubes is
subject to variation in most Cestodes, in the scolex as well as in the
segments. Indeed, even the lumen of the four longitudinal tubes
does not remain equal, as the dorsal or external tubes are more fully
developed and become thicker, whereas the ventral or internal ones
remain thin, and in some species quite disappear in the older
segments (figs. 185, 187). Moreover, very frequently connections are
established between the right and left longitudinal branches, as in
the head, where a “frontal anastomosis” develops, which in the
Tæniidæ usually takes the form of a ring encircling the rostellum
(fig. 190), and in the segments of a transverse anastomosis at each
posterior border, especially between the larger branches, and more
rarely between the smaller collecting tubes also (fig. 191).

Fig. 189.—Young Fig. 190.—Scolex of

Acantho­both­rium coro­‐ a cysti­cer­coid from
natum, v. Ben., with the Arion sp., with the
ex­cre­tory vessels out­‐ ex­cre­tory vessels
lined. Slightly enlarged. out­lined. (After
(After Pintner.) Pintner.)
The so-called “island” formation is another modification, i.e., at any
spot a vessel may divide and after a longer or shorter course the two
branches reunite, and this may appear in the collecting tubes
themselves as well as in their anastomoses. The above-mentioned
ring in the frontal commissure of the Tæniidæ is such an island;
similar rings also frequently encircle the suckers (fig. 190). In
extreme cases (Triænophorus, Ligula, Dibothriocephalus, etc.) this
island formation extends to all the collecting tubes and their
anastomoses. Instead of two or four longitudinal canals only,
connected by transverse anastomoses at the posterior border of the
segments, there is an irregular network of vessels, situated in the
cortical layer, from which the longitudinal branches, having again
subdivided, can only be distinguished at intervals, and even then not
in their usual number.
The opening of the longitudinal branches at the posterior end
requires more accurate investigation; it is true that a single terminal
bladder is mentioned as being present in many species, but this is
also disputed; when the original end proglottis has been cast off, the
longitudinal branches discharge separately. Some species possess
the so-called foramina secundaria, which serve as outlets for the
collecting tubes; they are generally at the neck, but may be situated
on the segments.
The contents of the excretory vessels is a clear fluid, the
regurgitation of which is prevented by the valves present at the
points of origin of the transverse anastomoses. The fluid contains in
solution a substance similar to guanine and xanthine.
Genital Organs.—With the exception of one genus (Diœcocestus,
Fuhrm.), in which the species are sexually differentiated, all the
Cestodes are hermaphroditic; the genitalia develop gradually in the
segments (never in the scolex), the male organs, as is usual in
hermaphroditic animals, forming earlier than the female. The
youngest proglottids generally do not exhibit even traces of
genitalia: these, as a rule, develop first in the older segments, and
the development proceeds onwards from segment to segment. In a
few exceptional cases (Ligula) the sexual organs are already
developed in the larval stage, but are only functional after the entry
of the parasite into the final host.

Fig. 191.—Proglottis of Tænia saginata, Goeze, showing genitalia.

C., transverse excretory canal; N., lateral longitudinal nerve; W.,
longitudinal excretory canal; T., testicles scattered throughout the
proglottis; Ut., opposite the central uterine stem (a closed sac); Ss.,
genital pore leading into the genital sinus; above the cirrus and
coiled vas deferens (V.d.), below the vagina (Vag.), bearing near its
termination a dilatation, the seminal receptacle; Vsc., the triangular
vitellarium, and above it (Shg.) the shell gland; leading from this to
the uterus is seen the short uterine canal, on either side of this the
two lobes of the ovary (Ov.). 10/1.

With the exception of the end portions of the vagina, cirrus and
uterus, all the parts of the genital apparatus lie in the medullary
layer, except only the vitellaria, which in many species are in the
cortical layer. The male apparatus consists of the testes, of which, as
a rule, there are a large number, 278 and which lie dorsal to the
median plane (fig. 185, T.); a vas efferens arises from each testis,
unites with contiguous vasa, and finally discharges into the muscular
vas deferens that is situated in about the middle of the segment.
According to the position of the genital pore, the vas deferens opens
on the lateral margin or in the middle line in the front of the
segment; it is much convoluted or twisted, and frequently possesses
a dilatation termed the vesicula seminalis. It finally enters the cirrus
pouch, which is usually elongated; within the cirrus pouch lies the
protrusible cirrus, which is not uncommonly provided with hooklets.
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