Geographic Range
Echinococcus multilocularis
lives in various regions around the world. In Europe, the endemic area includes
central and eastern France, Switzerland, Austria, and Germany. In Asia,
E. multilocularis
ranges from the zone of tundra from the White Sea to the Bering Strait and parts
of the former Soviet Union. The southern distribution in Asia extends from Turkey
eastwards through Afghanistan to the northern areas of Japan. In North America, this
cestode appears in subarctic Alaska and Canada as well as regions in north-central
and southern United States. (Gottstein 1992)
- Biogeographic Regions
- nearctic
- palearctic
- oriental
Habitat
The definitive hosts of
E. multilocularis
are usually
carnivores
.
Dogs
and
cats
are good definitive hosts.
Voles, muskrat, lemmings, Northern mole-vole
,
deer mouse
,
gerbils
, and striped
hamster
are typical intermediate hosts of this species, while ground beetles may act as paratenic
hosts.
Humans
may act as intermediate hosts as well. The habitat for this species is mainly sylvatic
(forest), and rural areas, rather than urban. However, once a hydatid cyst is brought
into a domestic community via dogs, humans, or farm animals, the
E. multilocularis
may begin a domestic cycle. Hunters and trappers tend to be the best human hosts.
Prevalence and distribution of
E. multilocularis
may vary with seasonal fluctuations (especially temperature fluctuations). Their
life span is inversely proportional to temperature. Once worms are in the host, they
reside in the small intestine, and travel to various other organs through the lymphatic
system. (Leiby and Nickel 1968), (Gottstein 1992), (Roberts and Janovy, Jr. 2000),
(Polydorou 1992)
- Habitat Regions
- temperate
- tropical
- terrestrial
- Terrestrial Biomes
- taiga
- savanna or grassland
- forest
- Other Habitat Features
- urban
- suburban
- agricultural
Physical Description
An adult E. multilocularis is 1.5 to 3.5 mm in length. It consists of a head, or scolex, a neck, and a segmented body, the strobila. This is the basic tapeworm anatomy. The rostellum is located on the frontal end surface, or apex, of the scolex. It has twenty-eight to thirty hooks each consisting of a shaft, root process, and blade. There are four suckers on the dorsolateral surface of the scolex, also called the rostellum. Like other tapeworms, this species has no mouth; they absorb nutrients through their external covering and microvilli. They also have no anus. The scolex is attached to the strobila via a long, thin neck. The segmented strobila consists of a number of reproductive sets called proglottids. Each proglottid is a complete set of reproductive organs, normally both male and female organs.
The larval stages differ from the adult. One distinct stage is the hydatid, a complex
cysticercus, developing from the hexacanth. They have an alveolar structure, ovoid
shaped, and make up a porous, spongy mass of daughter hydatids and protoscolices.
A protoscolex is juvenile scolex budded within a hydatid metacestode. They are a
grayish-white color with gelatinous contents and liters of fluids. Hydatids range
from 200 to 2000 micrometers. (Polydorou 1992), (Roberts and Janovy, Jr. 2000)
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
Development
The adult worm releases its proglottids filled with eggs from the definitive host
via the feces. Definitive hosts are carnivorous
mammals
, such as canines. The eggs contain the hexacanth, the "six-hooked larva", produced
through a process called embryogenesis. They require a short period of maturation
time after ingestion by the first intermediate host. The first intermediate host
may be an invertebrate or a vertebrate, usually rodents.
Humans
may also act as intermediate hosts. Once inside the intermediate host, the hexacanth
migrates through the gut wall to a parenteral or extraintestinal site within it.
At this time, the larva metamorphoses to a juvenile, or metacestode, with an introverted
and invaginated scolex and multiplies by endogenous and exogenous budding (asexual).
In
E. multilocularis
, the metacestode is a multilocular or alveolar hydatid. The germinal layer buds off
daughter hydatids and protoscolices, the stage ingested by the definitive host. This
part of the life cycle may take place in two to four months. Hydatids and protoscolices
will differentiate further to reach full adulthood once inside the definitive host.
The four stages involved in adult maturation are proglottisation, maturation, growth,
and segmentation. Proglottisation and maturation form the reproductive units. Growth
and segmentation lengthen the body. (Roberts and Janovy, Jr. 2000), (Thompson 1986),
(Gottstein 1992), (Morris and Richards 1992)
Reproduction
Adults of
E. multilocularis
are monoecious, they possess all parts of both male and female systems and reproduce
by self-fertilization. However, cross-insemination may occur between individuals.
They have both sexual and asexual life stages. Each segment has a set of male and
female systems. The reproductive life cycle typically requires two hosts, an intermediate
and a definitive host in which it will reach sexual maturity. The adult worm releases
its proglottids filled with eggs from the definitive host via the feces. (Roberts
and Janovy, Jr. 2000), (Thompson 1986), (Gottstein 1992), (Morris and Richards 1992)
- Parental Investment
- no parental involvement
Behavior
Echinococcosis multiocularis
activity after ingestion by definitive host varies. The protoscolices tend to be
quite active. This activity is a prerequisite to its establishment within the definitive
host. Once
E. multiocularis
has found its way into the small intestine, specifically in the crypts of the Lieberkuehn,
it remains there in a more sessile state. The organism inserts its rostellum into
the intestinal wall using its hooks and together with the suckers attaches to the
intestinal wall. A secretion from the rostellum may also anchor this species to the
host. There is little information about sensory mechanisms in
E. multiocularis
. However, the scolex is equipped with several sensory endings on its anterior surface
for both physical and chemical stimuli. Sensing its environment may allow the correct
placement of the scolex and the strobila on the intestinal surface. Dipersal of this
species, as mentioned above, varies depending on the definitive host. Humans acquire
the infection via ingestion of eggs from dog feces, and carnivores may acquire the
hydatid by consuming the intermediate host. (Roberts and Janovy, Jr. 2000), (Thompson
1986)
Communication and Perception
Cestodes
in general have sensory organs in the scolex, which are attached to longitudinal
nerves extending down the body. The nerves are attached to organs and the cestodes
can detect tactile stimulation.
- Communication Channels
- tactile
- Perception Channels
- tactile
Food Habits
Echinococcus multilocularis
feeds through the tegument, an external covering. Actual mechanisms of food absorption
include active transport, mediated diffusion, and simple diffusion. During the initial
stages of feeding, the adult tapeworm attaches to the epithelium of the small intestine
via the rostellar hooks and the suckers adhere to the small hairs or villi of the
intestine. These organs secure the parasite to the host. Microvilli called microtriches
layer the scolex and strobila. These mictotriches increase the absorptive area of
the tegument. Glycocalyx is found on the surface of microtriches. When glycocalyx
interacts with certain molecules some reactions may occur such as the inhibition of
host trypsin and the absorption of bile salts. Trypsin inhibition may protect the
organism against digestion by host enzymes and maintain the integrity of the surface
membrane. The rostellar glands aid nutrient absorption by releasing bioactive molecules
that process nutritive components of the host. The most important nutrient molecule
for energy metabolism is glucose. The only carbohydrates that can be metabolized
are glucose and galactose; they are both actively transported. Specifically, glucose
absorption in
E. multilocularis
is coupled to a sodium pump concentration. Amino acids are absorbed by active transport
while purines and pyrimidines are absorbed by facilitated diffusion. Lipids are also
absorbed by diffusion mechanisms. Although vitamins are necessary for the tapeworm,
mechanisms of absorption are unknown. Juvenile
E. multiocularis
absorb glucose in a similar fashion. (Roberts and Janovy, Jr. 2000)
- Primary Diet
-
carnivore
- eats body fluids
- Animal Foods
- body fluids
Predation
These animals are probably not preyed on directly but are ingested. Egg and larval mortality are high due to the parasite not reaching appropriate hosts
Ecosystem Roles
The definitive hosts of E. multilocularis are usually carnivores . Dogs and cats are good definitive hosts. Voles, muskrat, lemmings, Northern mole-vole , deer mouse , gerbils , and striped hamster are typical intermediate hosts of this species, while ground beetles may act as paratenic hosts. Humans may act as intermediate hosts as well.
- Ecosystem Impact
- parasite
- carnivorous mammals, Carnivora
- dogs, Canis lupus familiaris
- cats, Felidae
- voles, muskrat, lemmings, Northern mole-vole Arvicolinae
- gerbils, Gerbillinae
- striped hamster, Cricetinae
- humans, Homo sapiens
Economic Importance for Humans: Positive
There is no known positive economic importance for this species.
Economic Importance for Humans: Negative
Humans
can be parasitized by
E. multilocularis
. One of its own life stages, the hydatid cysts, is harmful to human populations.
The adult stages are tolerated by definitive hosts. Human infection usually occurs
because of close contacts with
dogs
. Eggs may be picked up from the dog's coat and ingestion from dog feces. The eggs
may be air borne due to the wind. Cultural or social practices determine the risk
to which individuals are exposed. Some of these cultural factors are: ethnicity,
occupational factors, economic variables, and husbandry or farming practices. In
humans, the greatest somatic site for infestation is the liver followed by pulmonary
(lung) infestation. The skeletal and nervous systems may also be affected. However,
the liver is almost always involved. There are no specific symptoms related to hydatid
disease-they will depend on what organ is affected by the disease. Once ingested
by humans, the cysts grow slowly and may continue for many years. Alveolar echinococcosis
of the liver can be progressive and metastasize to other organs. In humans, mortality
from toxic shock occurs if the cysts rupture. The processes by which the cestode
proliferates within the human have been described above (see reproduction and development).
(Uchino and Sato 1993), (Roberts and Janovy, Jr. 2000), (Thompson 1986), (Polydorou
1992)
- Negative Impacts
- injures humans
- causes or carries domestic animal disease
Conservation Status
There is no conservation status for E. multilocularis .
Other Comments
Echinococcus multilocularis
can have serious effects on animal populations, such as mortality in
humans
and farm animals from hydatid cyst infection. Several types of control and prevention
programs have been initiated. Animal control is one preventative measure, such as
reducing the number of stray or wild dogs. Legislation on dog control would also
prevent the spread of hydatid disease. In some areas infested by
E. multilocularis
, like Cyprus, local farmers who practice animal husbandry slaughter animals for meat
supplies. The potentially hydatid-infected carcasses of these animals are discarded
in unmanaged waste disposal sites. Some regulations on slaughtering practices are:
slaughtered animals may not be transported far and they must be inspected for worms
by a veterinarian. Drugs, such as prazquantel, kills hydatid cysts in canines. Mebendazole
and mitomycin C seem to inhibit hydatid cyst growth in humans. Once an infection
has occurred in humans, surgical management may ensue. Hepatic (liver) resection
is the only surgical therapy for
E. multilocularis
. This involves cutting away of hydatid lesions. Sometimes liver transplantation
must occur. (Morris and Richards 1992), (Thompson 1986), (Polydorou 1992), (Uchino
and Sato 1993)
Additional Links
Contributors
Renee Sherman Mulcrone (editor).
Christopher Bonadio (author), University of Michigan-Ann Arbor, Teresa Friedrich (editor), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- taiga
-
Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- urban
-
living in cities and large towns, landscapes dominated by human structures and activity.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- agricultural
-
living in landscapes dominated by human agriculture.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- asexual
-
reproduction that is not sexual; that is, reproduction that does not include recombining the genotypes of two parents
- parasite
-
an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- tactile
-
uses touch to communicate
- tactile
-
uses touch to communicate
- parasite
-
an organism that obtains nutrients from other organisms in a harmful way that doesn't cause immediate death
- causes disease in humans
-
an animal which directly causes disease in humans. For example, diseases caused by infection of filarial nematodes (elephantiasis and river blindness).
- causes or carries domestic animal disease
-
either directly causes, or indirectly transmits, a disease to a domestic animal
- carnivore
-
an animal that mainly eats meat
References
Brusca, R., G. Brusca. 2003. Invertebrates . Sunderland, Massachusetts: Sinauer Associates, Inc..
Gottstein, B. 1992. Echinococcus multilocularis infection: immunology and immundiagnosis. Advances in Parasitology , 31: 321-380.
Kahlill, L., A. Jones, R. Bray. 1994. Keys to the cestode parasites of vertebrates . Wallingford: CAB International.
Leiby, P., P. Nickel. 1968. Studies on the Sylvatic Echinococcosis: Ground Beetle Transmisson of Echinococcus multilocularis Leuckart, 1863, to Deer Mice, Peromyscus maniculatus (Wagner). Journal of Parasitology , 54: 536-537.
Morris, D., K. Richards. 1992. Hydatid Disease: Current Medical & Surgical Management . Butterworth-Heinemann, Ltd.: Oxford.
Polydorou, K. 1992. Echinococcosis Hydatidosis: The Problem and its Control, Case Study: Cyprus . Cyprus: Polydorou.
Roberts, L., J. Janovy Jr.. 2000. Foundations of Parasitology (6th ed.) . New York: McGraw-Hill.
Thompson, C. 1986. The Biology of Echinococcus and Hydatid Disease . London: George Allen & Unwin.
Uchino, J., N. Sato. 1993. Alveolar Echinococcosis of the Liver . Sapporo: Kokoku Printing Co. Ltd..