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Fasciola hepatica
Fasciola hepatica, also known as the common liver fluke or sheep
Fasciola hepatica
liver fluke, is a parasitic trematode (fluke or flatworm, a type of
helminth) of the class Trematoda, phylum Platyhelminthes. It infects the
livers of various mammals, including humans. The disease caused by the
fluke is called fasciolosis or fascioliasis, which is a type of helminthiasis
and has been classified as a neglected tropical disease.[2] Fasciolosis is
currently classified as a plant/food-borne trematode infection, often
acquired through eating the parasite's metacercariae encysted on plants.[3]
F. hepatica, which is distributed worldwide, has been known as an
important parasite of sheep and cattle for decades and causes significant
economic losses in these livestock species, up to £23 million in the UK Adult Fasciola hepatica specimen
alone.[4] Because of its relatively large size and economic importance, it
Scientific classification
has been the subject of many scientific investigations and may be the best-
known of any trematode species. F. hepatica's closest relative is Fasciola Kingdom: Animalia
gigantica. These two flukes are sister species; they share many Phylum: Platyhelminthes
morphological features and can mate with each other.[5]
Class: Trematoda
Order: Plagiorchiida
Family: Fasciolidae
Contents
Genus: Fasciola
Life cycle
Species: F. hepatica
Morphology and anatomy
Tegument Binomial name
Digestive system
Fasciola hepatica
Respiratory system
Excretory system Linnaeus, 1758[1]
Nervous system and sensory organs
Reproductive system
Prevalence
Parasitic adaptations
Epidemiology
Fasciolosis
Diagnosis
See also
References
External links
Life cycle
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Fasciola hepatica occurs in the liver of a definitive host and its lifecycle is
indirect. Definitive hosts of the fluke are cattle, sheep, and buffaloes. Wild
ruminants and other mammals, including humans, can act as definitive hosts
as well.[6] The life cycle of F. hepatica goes through the intermediate host and
several environmental larval stages.[7] Intermediate hosts of F. hepatica are
air-breathing freshwater snails from the family Lymnaeidae. Although
several lymnaeid species susceptible to F. hepatica have been described, the
parasite develops only in one or two major species on each continent. Galba
truncatula is the main snail host in Europe, partly in Asia, Africa, and South
America. Lymnaea viator, L. neotropica, Pseudosuccinea columella, and L.
cubensis are most common intermediate hosts in Central and South
America.[5][6] Several other lymnaeid snails may be naturally or
experimentally infected with F. hepatica, but their role in transmission of the
fluke is low.[5] The list of lymnaeid snails that may serve as natural or
experimental intermediate hosts of F. hepatica include:[8]
Austropeplea ollula
Austropeplea tomentosa
Austropeplea viridis
Fossaria bulimoides
Galba truncatula
Lymnaea cousini Galba truncatula, an amphibious
Lymnaea cubensis freshwater lymnaeid snail that
Lymnaea diaphana serves as the main intermediate
Lymnaea humilis host of Fasciola hepatica in Europe
Lymnaea neotropica
Lymnaea occulta
Lymnaea stagnalis
Lymnaea viatrix
Omphiscola glabra
Pseudosuccinea columella
Radix auricularia
Radix lagotis
Radix natalensis
Radix peregra
Radix rubiginosa
Stagnicola caperata The lifecycle of Fasciola hepatica
Stagnicola fuscus
Stagnicola palustris
Stagnicola turricula
The metacercariae are released from the freshwater snail as cercariae, and form cysts on various surfaces including
aquatic vegetation. The mammalian host then eats this vegetation and can become infected. Humans can often acquire
these infections through eating freshwater plants such as watercress. Inside the duodenum of the mammalian host, the
metacercariae are released from within their cysts. From the duodenum, they burrow through the lining of the intestine
and into the peritoneal cavity. They then migrate through the intestines and liver, and into the bile ducts. Inside the bile
ducts, they develop into an adult fluke.[9] In humans, the time taken for F. hepatica to mature from metacercariae into
an adult fluke is roughly 3 to 4 months. The adult flukes can then produce up to 25,000 eggs per fluke per day.[10] These
eggs are passed out via stools and into freshwater. Once in freshwater, the eggs become embryonated, allowing them to
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hatch as miracidia, which then find a suitable intermediate snail host of the Lymnaeidae family. Inside this snail, the
miracidia develop into sporocysts, then to rediae, then to cercariae. The cercariae are released from the snail to form
metacercariae and the life cycle begins again.[9]
Tegument
The outer surface of the fluke is called the tegument. This is composed of
scleroprotein, and its primary function is to protect the fluke from the
destructive digestive system of the host.[12] Its also used for renewal of the
surface plasma membrane and the active uptake of nutrients.[13] On the
surface of the tegument are also small spines. Initially, these spines are
single-pointed, then, just prior to the fluke entering the bile ducts, they A simple diagram to show the
become multipointed. At the anterior end of the fluke, the spines have difference between the teguments of
between 10 and 15 points, whereas at the posterior end, they have up to 30 free-living and parasitic flatworms:
points.[14] The tegument is a syncytial epithelium. This means it is made a. shows the syncytial epithelial
from the fusion of many cells, each containing one nucleus, to produce a tegument found in parasitic
flatworms, such as F. hepatica. b.
multinucleated cell membrane. In the case of F. hepatica, no nuclei are in
shows the multicellular,
the outer cytoplasm between the basal and apical membranes. Thus, this nonsyncytial, epithelia, found in
region is referred to as anucleate. Instead, the nuclei are found in the cell nonparasitic, free-living flatworms.
bodies, also known as tegumental cells, these connect to the outer cytoplasm
via thin cytoplasmic strands. The tegumental cells contain the usual
cytoplasmic organelles (mitochondria, Golgi bodies, and endoplasmic reticulum).[15] The tegument plays a key role in
the fluke’s infection of the host. Studies have shown that certain parts of the tegument (in this case, the antigen named
Teg) can actually suppress the immune response of the mammalian host. This means that the fluke is able to weaken
the immune response, and increase its chances of a successful infection. A successful infection is needed for the fluke to
have enough time to develop into an adult and continue its lifecycle.[16]
Digestive system
The alimentary canal of F. hepatica has a single mouth which leads into the blind gut; it has no anus. The mouth is
located within the anterior sucker on the ventral side of the fluke. This mouth leads to the pharynx, which is then
followed by a narrow oesophagus. The oesophagus, which is lined with a thin layer of epithelial cells, then opens up into
the large intestine. As no anus is present, the intestine branches, with each branch ending blindly near the posterior end
of the body.[17] Fflukes migrate into smaller capillaries and bile ducts when feeding within the host. They use their
mouth suckers to pull off and suck up food, bile, lymph, and tissue pieces from the walls of the bile ducts.[17] F. hepatica
relies on extracellular digestion which occurs within the intestine of the host. The waste materials are egested through
the mouth. The nonwaste matter is adsorbed back in through the tegument and the general surface of the fluke. The
tegument facilitates this adsorption by containing many small folds to increase the surface area.[17]
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Respiratory system
F. hepatica has no respiratory organs: the adult flukes respire anaerobically
(without oxygen). Glycogen taken from within the host is broken down by
glycolysis to produce carbon dioxide and fatty acids. This process provides
the fluke with energy.[18] In contrast, the free-living miracidia stages of the
parasite generally develop within oxygen-rich environments. The free-living
stages of the parasite are thought to respire aerobically, to gain the most
energy from their environment.[19]
Excretory system
F. hepatica's excretory system contains a network of tubules surrounding
one main excretory canal. This canal leads to the excretory pore at the
posterior end of the fluke. This main canal branches into four sections
within the dorsal and ventral regions of the body. The role of F. hepatica's
excretory system is excretion and osmoregulation.[18] Each tubule within the
excretory system is connected to a flame cell, otherwise known as
protonephridia. These cells are modified parenchyme cells. In F. hepatica,
their role is to perform excretion, but more importantly, osmoregulatory
functions. Flame cells are therefore primarily used to remove excess Image showing the location of the
water.[18] mouth, labelled mo, and the anterior
sucker, as labelled sckr
Reproductive system
F. hepatica adult flukes are hermaphrodite; each contains both male and female reproductive organs. The male and
female reproductive organs open up into the same chamber within the body, which is called the genital atrium. The
genital atrium is an ectodermal sac which opens up to the outside of the fluke via a genital pore.[20] The testes are
formed of two branched tubules, these are located in the middle and posterior regions of the body. From the epithelium
lining of the tubules, sperm is produced. The sperm then passes into the vas deferens and then into the seminal vesicle.
From the seminal vesicle projects the ejaculatory duct, and this opens into the genital atrium, and many prostate glands
surround this opening.[20] The right side of the anterior testis has a branched, tubular ovary. From here, a short oviduct
passes to the vitelline duct. This duct connects, via a junction, the ovaries, the uterus, and the yolk reservoir. From this
junction, the uterus opens into the genital atrium; this opening is surrounded by Mehlis glands. In some flukes, the
terminal end of the uterus is strengthened with muscles and spines.[20]
F. hepatica reproduces both sexually, via the hermaphrodite adult flukes, and asexually. The miracidia can reproduce
asexually within the intermediate snail host.[22]
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Prevalence
Currently, F. hepatica has one of
the widest geographical spread
of any parasitic and vector-borne
disease. Originating in Europe, it
has expanded to colonize over 50
Fasciola hepatica prevalence. The countries, covering all continents
countries in red are those with high except Antarctica.[23] In contrast,
prevalence, those in orange have F. gigantica is generally
low-medium prevalence.[23][24][25][26] considered more geographically
restricted to the tropical regions
of Africa, Asia, and the Middle
East, with some overlap between the two species.[24]
Climate affects both F. hepatica and its definitive host, the snail. For
example, the development of F. hepatica miracidia and larvae, and the
reproduction of Galba truncatula, require a temperature range of 10 to Diagram of the main organ systems
25°C. In addition, they both require high levels of moisture in the air, as of F. hepatica throughout the
both are at risk of desiccation. Due to this, the prevalence, along with the progressive life stages of the fluke
(1938). A - egg; B - miracidium; C -
intensity of infection, of F. hepatica is primarily dependent on rainfall levels
sporocyst; D - rediae, E - immature
and temperature.[23]
cercaria, F - cercaria, G - encysted
stage, H - adult fluke (nervous and
Parasitic adaptations reproductive systems omitted)
Epidemiology
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For more information on the epidemiology – see the disease page, fasciolosis
Infection begins when cyst-covered aquatic vegetation is eaten or when water containing metacercariae is drunk. In the
United Kingdom, F. hepatica frequently causes disease in ruminants, most commonly between March and
December.[30]
Humans become infected by eating watercress or by drinking 'Emoliente', a Peruvian drink that uses drops of
watercress juice. Cattle and sheep are infected when they consume the infectious stage of the parasite from low-lying,
marshy pasture.[30]
Human infections have been reported from more than 75 countries around the world. In Asia and Africa, people are
infected both by F. hepatica and F. gigantica whereas human fasciolosis is caused only by F. hepatica in South and
Central America and Europe.[31]
The presence of F. hepatica can interfere with the detection of bovine tuberculosis in cattle. Cattle co-infected with F.
hepatica, compared to those infected with M. bovis alone, react weakly to the single intradermal comparative cervical
tuberculin (SICCT) test (http://ahvla.defra.gov.uk/External_OV_Instructions/TB_Instructions/Skin_Test/index.htm).
Therefore, an infection from F. hepatica can make it difficult to detect bovine tuberculosis, this is, of course, a major
problem in the farming industry.[32]
Fasciolosis
Both F. hepatica and F. gigantica can cause fasciolosis. Human
symptoms vary depending on whether the disease is chronic or acute.
During the acute phase, the immature worms begin penetrating the
gut, causing symptoms of fever, nausea, swollen liver, skin rashes, and
extreme abdominal pain.[33] The chronic phase occurs when the worms
mature in the bile duct, and can cause symptoms of intermittent pain,
jaundice, and anemia.[33] In cattle and sheep, classic signs of fasciolosis Slide showing Fasciola hepatica's
internal organs
include persistent diarrhea, chronic weight loss, anemia, and reduced
milk production.[34] Some remain asymptomatic. F. hepatica can cause
sudden death in both sheep and cattle, due to internal hemorrhaging and liver damage.[4]
Fasciolosis is an important cause of both production and economic losses in the dairy and meat industries. Over the
years, the prevalence has increased and it is likely to continue increasing in the future.[35] Livestock are often treated
with flukicides, which are chemicals toxic to flukes. The two chemicals used are triclabendazole and bithionol.
Ivermectin, which is widely used for many helminthic parasites, has low effectivity against F. hepatica, as does
praziquantel.[36][37] For humans, the type of control depends on the setting. One important method is through the strict
control over the growth and sales of edible water plants such as watercress. This is particularly important in highly
endemic areas. Some farms are irrigated with polluted water, hence, vegetables farmed from such land should be
thoroughly washed and cooked before being eaten.[9]
The best way to prevent fasciolosis is by reducing the lymnaeid snail population or separating livestock from areas with
these snails.[34] These two methods are not always the most practical, so control by treating the herd before they are
potentially infected is commonly practiced.
Diagnosis
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See also
List of parasites (human)
Veterinary parasitology
Trematoda
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External links
University of Michigan Animal Diversity Web (http://animaldiversity.ummz.umich.edu/accounts/Fasciola_hepatica/)
Parasite Fasciola(parasite.org.au) (http://parasite.org.au/para-site/text/fasciola-text.html)
Stanford University Fascioliasis Info Page (http://www.stanford.edu/class/humbio103/ParaSites2001/fascioliasis/Fa
sciola.htm)
Encyclopedia of Life (http://eol.org/pages/590853/hierarchy_entries/51054252/overview)
Taxonomy and nomenclature at ITIS.gov (https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&sear
ch_value=56135)
Molecular database at UniProt (http://www.uniprot.org/uniprot/P80342)
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