Classification:
Taxonomic ranks under review (cf. Encyclopedic Reference of Parasitology,
2001, Springer-Verlag)
Metazoa (Animalia) (multicellular eukaryotes, animals)
Platythelminthes (flatworms)
Cestoda (tapeworms)
Eucestoda (segmented, hermaphroditic)
Cyclophyllidea (terrestrial cycles, scolex with suckers)
Family:
Taeniidae
Cyclophyllidean
tape-worms have flat ribbon-like bodies, with an anterior scolex (hold-fast
organ with suckers and sometimes hooks) and a posterior tape (strobila)
made up of segments (proglottids). Adult worms lack a gut (they absorb
nutrients) and they are hermaphroditic (segments containing both male
and female reproductive organs). They have indirect life-cycles involving
encystment of larvae (metacestodes) in the tissues of intermediate hosts
and their transmission to definitive hosts by carnivorism. Various species
are parasitic in mammals, birds, reptiles and amphibians. Adult stages
are rarely pathogenic, but the encysted larval stages may cause serious
space-occupying lesions, including hydatid cysts in humans.
Echinococcus
granulosus [this species causes hydatid
disease in mammals, including humans]
Parasite
morphology:
Tape-worms form three different developmental stages: eggs; larvae;
and adults. Adult E. granulosus worms are small (2-6mm long) and have
a scolex with only three attached segments. The scolex has four lateral
suckers and the rostellum is non-retractable and armed with a double
crown of 28-50 recurved hooks. The anterior segment is immature, the
middle segment is mature with functional testes and ovaries, and the
posterior segment is gravid with the uterus filled with eggs. The eggs
are typical for most taeniid species and are small and round (30-43ยตm
in diameter), thick-shelled and contain a hexacanth (6-hooked) embryo
(oncosphere). The encysted larval (metacestode) stage is known as a
bladder-worm or hydatid, and it produces multiple infective stages (protoscoleces,
apparent as invaginated scolices already containing suckers and hooks)
either directly from the germinal layer of the cyst wall, or by forming
brood sacs (hydatid sand) by endogenous (internal) or exogenous (external)
budding of the germinal layer. E. granulosus forms fluid-filled
unilocular cysts with endogenous budding of brood capsules, E. vogeli
forms fluid-filled polycystic cysts with exogenous budding, and E.
multilocularis forms fluid-free multilocular or alveolar cysts
with exogenous budding.
Host range: E. granulosus
occurs in most sheep and cattle producing areas around the world, being
most prevalent in South America, East Africa, Southeast Asia and China.
Canids (dogs, dingoes, wolves, and coyotes) act as definitive hosts
for adult worms, while omnivorous/herbivorous mammals (humans, domestic
animals and wildlife) serve as intermediate hosts for encysted larval
stages.
Parasite species |
Definitive
host |
Intermediate
host |
Metacestode |
Cyst
morphology |
Echinococcus
granulosus |
candid |
omnivore |
unilocular
hydatid cyst |
fluid-filled sphere with germinal membrane proliferating endogenously
to form brood capsules |
Echinococcus
vogeli |
bush
dog |
paca/rat |
polycystic
hydatid cyst |
fluid-filled with germinal membrane budding exogenously to form
new cysts and endogenously to form septae |
Echinococcus
multilocularis |
dog/cat |
rodent |
multilocular
(alveolar) hydatid cyst |
no free fluid, germinal membrane budding exogenously to form multiple
cysts |
Site
of infection: The
small adult tape-worms attach to the mucosa of the small intestines in
dogs, sometimes in their thousands. The larval stages (hydatids) most
commonly infect visceral tissues and organs, especially the liver, in
their mammalian intermediate hosts, although cysts may be found in many
other locations, including the brain and long bones.
Pathogenesis:
The adult stages are considered benign and do not cause disease in dogs,
as the worms do not invade or feed on host tissues. Encysted larval stages
generally do not cause clinical disease in domestic livestock as they
are often confined to visceral tissues. However, significant pathological
changes occur in humans when the slowly-growing cysts put pressure on
surrounding tissues and produce chronic space-occupying lesions.Cysts
may grow around 1 mm per month and can become extremely large, up to 30cm
in diameter with litres of fluid containing thousands of protoscoleces.
Organ enlargement may be accompanied by a variety of clinical signs depending
on the size and location of the cysts. Compression of liver may result
in jaundice, portal hypertension and abdominal distention. Cysts in the
lung may cause haemoptysis (coughing up blood), dyspnoea (difficulty breathing)
and chest pain. Cysts in the brain or spinal cord can provoke acute inflammatory
responses and numerous neurological sequalae, including epilepsy and blindness.
Cyst rupture has been associated with acute clinical signs (such as peritonitis
and pneumothorax), and the sudden release of hydatid fluid may cause severe
allergic reactions (such as asthma and anaphylactic shock). Protoscoleces
released from ruptured cysts can regress and form new hydatid cysts throughout
the body.
Mode
of transmission:
Tape-worms have an indirect life-cycle involving predator-prey transmission
between definitive (canid) and intermediate (mammalian) hosts. Mature
tape-worms release numerous thick-shelled eggs which are excreted with
dog faeces. The eggs are very resistant to external environmental conditions
and can survive for months on pasture. Herbivores and omnivores become
infected by ingesting eggs; either on herbage, in water, or by hand-to-mouth
transfer. Following ingestion, the eggs hatch releasing the oncosphere
which uses its three pairs of hooks to penetrate the gut, enter the circulation
and settle in various organs and tissues (frequently in the liver after
being filtered out by portal capillaries). They form hydatid cysts over
many months and eventually produce multiple infective protoscoleces. When
mature cysts in offal or carcases are eaten by canids, the cyst wall is
digested away freeing the protoscoleces, which evaginate and attach to
the small intestinal mucosa. They mature to adult worms in about 8 weeks
and may live for 5-20 months. Various strains of E. granulosus
have been recognized based on differences in parasite morphology, development,
biochemistry, genetics and host specificity. Strains are often adapted
to particular intermediate host species and do not develop well in other
species. Infections are well adapted to pastoral cycles involving farm
dogs and domestic livestock (notably sheep and cattle), as well as sylvatic
cycles involving wild carnivores (wolves, coyotes, dingoes) and free-ranging
herbivores (such as deer, moose and wallabies). Infections in human populations
occur more frequently in rural areas, particularly where local traditions
are conducive to transmission; e.g. feeding dogs offal, eating dog intestines,
not burying the dead, and even using dog faeces to tan hides.
Differential
diagnosis:
Infections in dogs may
be diagnosed by the detection of eggs, and occasionally worms, in faecal
samples. Immuno-coprological tests have also been developed to detect
parasite antigens in faecal samples. Infections in intermediate hosts
are generally diagnosed well after the larvae have encysted. Clinical
symptoms of a slow-growing tumour accompanied by eosinophilia are suggestive.
Cysts may be visualized by various medical imaging techniques (computerized
axial tomography (CAT) scans, X-rays, ultrasound). Several immunoserological
tests have been developed to detect host antibodies against crude and
purified parasite antigens, and an intradermal (Casoni) test using hydatid
fluid has been used in surveys.
Treatment and control:
Despite some promising
indications, the treatment of hydatid disease with conventional anthelmintic
drugs has not proven wholly effective, being complicated by the large
size and inaccessible location of cysts and their thick, possibly impenetrable,
walls. Variable results have been obtained using praziquantel and mebendazole,
while albendazole and niclosamide have been less effective. The only remaining
treatment option is for the surgical removal of cysts, provided they are
in favourable sites. Surgeons must take care not to rupture cysts as protoscoleces
may spread to new sites to form more cysts. Scolicide chemicals, such
as cetrimide, may also be used during surgery to sterilize excision sites.
In contrast, infections by adult worms in dogs can be successfully treated
with praziquantel, and it is advisable to confine dogs and/or use purgatives
to facilitate the collection and disposal of infected faeces. Preventing
dogs from becoming infected involves eliminating offal and other potentially
infected material from their diets, curbing their hunting behaviour, properly
disposing of carcases in the field, and culling wild and feral dogs. Several
countries have developed highly successful hydatid eradication campaigns
based around dog management and treatment. Recently, a recombinant vaccine
has been developed to prevent hydatid formation in domestic herbivores,
and is undergoing further evaluation. While control may be possible in
situations involving pastoral cycles, there will be many problems accessing
wildlife involved in sylvatic cycles.
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