Echinococcus
 

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 (30-43µm 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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