Paramphistomum Cervi
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''Paramphistomum cervi'', the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of '' Paramphistomum'', is a
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The ent ...
flat worm belonging to the
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
Trematoda Trematoda is a class of flatworms known as trematodes, and commonly as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is a mol ...
. It is a tiny fluke mostly parasitising
livestock Livestock are the Domestication, domesticated animals that are raised in an Agriculture, agricultural setting to provide labour and produce diversified products for consumption such as meat, Egg as food, eggs, milk, fur, leather, and wool. The t ...
ruminants Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by Enteric fermentation, fermenting it in a specialized stomach prior to digestion, principa ...
, as well as some wild
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
. Uniquely, unlike most parasites, the adult worms are relatively harmless, but it is the developing juveniles that cause serious disease called paramphistomiasis (or classically amphistomosis), especially in
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
and
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
. Its
symptoms Signs and symptoms are diagnostic indications of an illness, injury, or condition. Signs are objective and externally observable; symptoms are a person's reported subjective experiences. A sign for example may be a higher or lower temperature ...
include profuse
diarrhoea Diarrhea (American English), also spelled diarrhoea or diarrhœa (British English), is the condition of having at least three loose, liquid, or watery bowel movements in a day. It often lasts for a few days and can result in dehydration d ...
,
anaemia Anemia (also spelt anaemia in British English) is a blood disorder in which the blood has a reduced ability to carry oxygen. This can be due to a lower than normal number of red blood cells, a reduction in the amount of hemoglobin availab ...
,
lethargy Lethargy is a state of tiredness, sleepiness, weariness, fatigue, sluggishness, or lack of energy. It can be accompanied by depression, decreased motivation, or apathy. Lethargy can be a normal response to inadequate sleep, overexertion, overw ...
, and often result in death if untreated. It can be found in many regions of the world, including
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
,
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
,
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, and
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
. It is most commonly found in
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
regions. The most debilitating cases are reported in Europe from
Bulgaria Bulgaria, officially the Republic of Bulgaria, is a country in Southeast Europe. It is situated on the eastern portion of the Balkans directly south of the Danube river and west of the Black Sea. Bulgaria is bordered by Greece and Turkey t ...
,
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
,
France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
, and
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
and also in Asia from
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
,
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
, and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The parasitic infection was first described from
Punjab Punjab (; ; also romanised as Panjāb or Panj-Āb) is a geopolitical, cultural, and historical region in South Asia. It is located in the northwestern part of the Indian subcontinent, comprising areas of modern-day eastern Pakistan and no ...
, India.


Description

Adult ''P. cervi'' is
conical In geometry, a cone is a three-dimensional figure that tapers smoothly from a flat base (typically a circle) to a point not contained in the base, called the ''apex'' or '' vertex''. A cone is formed by a set of line segments, half-lines, ...
in shape, the anterior end tapering and the posterior being broad, and pink in colour. The colour is due to its own
haemoglobin Hemoglobin (haemoglobin, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobi ...
. They are 5–13 mm long, 2–5 mm wide, with the ventral side somewhat concave while the dorsal side is convex. It has two suckers, an anterior oral sucker and a posterior larger ventral sucker, hence the
generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...
name (
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
: ''para'' meaning "besides", ''amphi'' meaning "on both sides", and ''stoma'' for "mouth"). The tegumental surface is highly corrugated with transverse folds alternating with grooves and is spineless, which is uncharacteristic of trematodes. The genital pore is situated at the anterior third of the body. There are two types of bulbous shaped sensory papillae on the surface, each measuring 10–15 μ in diameter at the base with nipple-like tips; one has a short
cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...
on top. Clusters of papillae on the ventral surface and around the anterior suckers are larger in size and number; while there are few on the dorsal surface. As hermaphrodite, both male and female
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are al ...
s are present towards the posterior region of the body.
Testes A testicle or testis ( testes) is the gonad in all male bilaterians, including humans, and is homologous to the ovary in females. Its primary functions are the production of sperm and the secretion of androgens, primarily testosterone. The ...
are slightly lobed and are located anterior to the
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
. Eggs are clear shell and measure about 140 x 80 μm; barrel-shaped with operculum at one end.


Life cycle

The
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from conception to reproduction *Life-cycle hypothesis, in economics *Erikson's stages of psy ...
is indirect, involving a ruminant as
definitive host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...
,
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
as
intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...
, and an interval of external phases in water and plants. The sexually mature
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
adult self-fertilises in the mammalian
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants. The rumen and the reticulum make up the reticulorumen in ruminant animals. The diverse microbial communities in the rumen allows it to serve as the primary si ...
, and retains the eggs inside its
uterus The uterus (from Latin ''uterus'', : uteri or uteruses) or womb () is the hollow organ, organ in the reproductive system of most female mammals, including humans, that accommodates the embryonic development, embryonic and prenatal development, f ...
. Eggs are then released in the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...
intestine and are expelled along with
faeces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
. Eggs are deposited in water and hatch at optimal temperature of in 12–17 days to become ciliated
miracidia The miracidium is the second stage in the life cycle of trematodes. When trematode eggs are laid and come into contact with fresh water, they hatch and release miracidium. In this phase, miracidia are ciliated and free-swimming. This stage is comp ...
. The non-feeding miracidia often swim through water searching for a suitable snail host, either until they reach the host or die. Most common intermediate hosts are snails belonging to the
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
''
Bulinus ''Bulinus'' is a genus of small tropical freshwater snails, aquatic gastropod mollusks in the family Bulinidae, the ramshorn snails and their allies.MolluscaBase eds. (2020). MolluscaBase. Bulinus O. F. Müller, 1781. Accessed through: World R ...
'', ''
Planorbis ''Planorbis'' is a genus of air-breathing freshwater snails, aquatic animal, aquatic pulmonate gastropod molluscs in the family (biology), family Planorbidae, the sheep, ram's horn snails, or planorbids. All species in this genus have sinistral o ...
'', ''
Physa ''Physa'' is a genus of small, left-handed or sinistral, air-breathing freshwater snails, aquatic pulmonate gastropod molluscs in the subfamily Physinae of the family Physidae.MolluscaBase eds. (2021). MolluscaBase. Physa Draparnaud, 1801. Acces ...
'', ''
Stagnicola ''Stagnicola'' can refer to: * ''Stagnicola'' (fungus), a fungus genus * ''Stagnicola'' (gastropod), a freshwater snail genus {{Genus disambiguation ...
'' and '' Pseudosuccinea''. When sensing the soft outer surface of a snail, they attach and burrow through the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
and into the tissue. Young snails are more susceptible to such penetration. In the snail tissue the miracidia lost their cilia within 12 hours and get transformed into sporocysts (~53 x 93 μm). By 11th day they develop up to 8 rediae, which are rapidly liberated. After 10 days the rediae mature and each contains 15–30
cercariae A cercaria (plural cercariae) is a larval form of the trematode class of parasites. It develops within the germinal cells of the sporocyst or redia. A cercaria has a tapering head with large penetration glands. It may or may not have a long swim ...
. The cercariae are released to grow further which may take about 13 days. Mature cercariae are characterised by two eyespots and a long slender tail. They are stimulated by light through the sensory eyespots in sunlight and leave the snail. They swim about in water until they find plants or other suitable substrata, to which they adhere and
encyst A microbial cyst is a resting or dormant stage of a microorganism, that can be thought of as a state of suspended animation in which the metabolic processes of the cell are slowed and the cell ceases all activities like feeding and locomotion. ...
to become
metacercariae Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two Sucker (zoology), suckers: one ventral and the other Mouth, oral. Trematodes are covered by a Tegument (helminth), tegument, that protects the o ...
. The process of this cyst formation requires about 20 minutes only. Metacercariae are infective larvae but cannot resist
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
, hence soon die out if suitable host is not found; but under constantly moist conditions, they can survive for up to 1 year and are capable of overwintering. The mammalian hosts harbour the infective larvae by
ingestion Ingestion is the consumption of a substance by an organism. In animals, it normally is accomplished by taking in a substance through the mouth into the gastrointestinal tract, such as through eating or drinking. In single-celled organisms, inge ...
. Once they reach the
duodenum The duodenum is the first section of the small intestine in most vertebrates, including mammals, reptiles, and birds. In mammals, it may be the principal site for iron absorption. The duodenum precedes the jejunum and ileum and is the shortest p ...
and
jejunum The jejunum is the second part of the small intestine in humans and most higher vertebrates, including mammals, reptiles, and birds. Its lining is specialized for the absorption by enterocytes of small nutrient molecules which have been pr ...
, their
cyst A cyst is a closed sac, having a distinct envelope and division compared with the nearby tissue. Hence, it is a cluster of cells that have grouped together to form a sac (like the manner in which water molecules group together to form a bubb ...
s are cast off. Excystment is influenced by changing physicochemical conditions (such as temperature, substance concentration, and pH) inside the
alimentary tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascular system. T ...
. Then they penetrate and feed on tissue of the gut wall of the small intestine. Once they develop sufficiently they migrate to the rumen, where the juveniles mature and produce
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and special episodes of a series made specially for release in home video formats without prior showings on television or in theaters, though the ...
. The period of rumen entry to egg laying takes about 60 to 120 days. However, the lifespan of an adult is unknown.


Pathogenicity and pathology

Paramphistomiasis causes enteritis and anaemia in livestocks mammals and result in substantial production and economic losses. Adults attach to the villi in the rumens of the hosts and sap nutrients from the intestine, although they can wander into the
bile Bile (from Latin ''bilis''), also known as gall, is a yellow-green/misty green fluid produced by the liver of most vertebrates that aids the digestion of lipids in the small intestine. In humans, bile is primarily composed of water, is pro ...
and
pancreatic The pancreas (plural pancreases, or pancreata) is an organ of the digestive system and endocrine system of vertebrates. In humans, it is located in the abdomen behind the stomach and functions as a gland. The pancreas is a mixed or heterocrine ...
ducts, as do other trematodes. Pathological symptoms are produced by immature flukes. When the young flukes start to gather in the intestine, there is a watery and fetid diarrhoea which is often associated with high mortality (even up to 80–90%) in ruminants. At a given time, as many as 30,000 flukes may accumulate, fervently attacking the duodenal mucosa to induce acute enteritis. Surprisingly, the adult flukes are regarded as
commensal Commensalism is a long-term biological interaction (symbiosis) in which members of one species gain benefits while those of the other species neither benefit nor are harmed. This is in contrast with mutualism, in which both organisms benefit f ...
s and non-pathogenic. However, they do cause the intestinal villi to erode and instil
inflammation Inflammation (from ) is part of the biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or irritants. The five cardinal signs are heat, pain, redness, swelling, and loss of function (Latin ''calor'', '' ...
. Liver tissue are generally damaged extensively, indicated by swelling,
haemorrhage Bleeding, hemorrhage, haemorrhage or blood loss, is blood escaping from the circulatory system from damaged blood vessels. Bleeding can occur internally, or externally either through a natural opening such as the mouth, nose, ear, urethra, vag ...
, discolouration,
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
, bile duct
hyperplasia Hyperplasia (from ancient Greek ὑπέρ ''huper'' 'over' + πλάσις ''plasis'' 'formation'), or hypergenesis, is an enlargement of an organ or tissue caused by an increase in the amount of Tissue (biology), organic tissue that results from ...
, and
fibrosis Fibrosis, also known as fibrotic scarring, is the development of fibrous connective tissue in response to an injury. Fibrosis can be a normal connective tissue deposition or excessive tissue deposition caused by a disease. Repeated injuries, ch ...
. Paramphistomiasis is responsible for severe economic losses to
milk Milk is a white liquid food produced by the mammary glands of lactating mammals. It is the primary source of nutrition for young mammals (including breastfeeding, breastfed human infants) before they are able to digestion, digest solid food. ...
,
meat Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
and
wool Wool is the textile fiber obtained from sheep and other mammals, especially goats, rabbits, and camelids. The term may also refer to inorganic materials, such as mineral wool and glass wool, that have some properties similar to animal w ...
production since the flukes take nutrients from their hosts, which leads to weight loss and physiological decline.


Diagnosis and treatment

Symptoms are usually visible on the behaviour of the host. Infected sheep and cattle become severely
anorexic Anorexia nervosa (AN), often referred to simply as anorexia, is an eating disorder characterized by Calorie restriction, food restriction, body image disturbance, fear of gaining weight, and an overpowering desire to be thin. Individuals wit ...
or digest food inefficiently and become unthrifty. Continuous diarrhoea is an obvious indication of heavy infection in the digestive system, thus a primary diagnosis. The fluid faeces are examined to identify immature flukes. Paramphistomiasis is considered a neglected tropical disease, with no prescription drug for treatment and control. Thus management of infection is based mainly on to reduce the host snail population. There are some treatment strategies demonstrated. A common regime is to drench with hexachloroethane-bentonite-water suspension, which is highly effective against adult parasites. Treatments with reported success (efficacies >90%) are resorantel,
oxyclozanide Oxyclozanide is a salicylanilide anthelmintic. It is used in the treatment and control of fascioliasis in ruminants, mainly domestic animals such as cattle, sheep, and goats. It mainly acts by uncoupling of oxidative phosphorylation in flukes. Al ...
, clorsulon,
ivermectin Ivermectin is an antiparasitic drug. After its discovery in 1975, its first uses were in veterinary medicine to prevent and treat heartworm and acariasis. Approved for human use in 1987, it is used to treat infestations including head lice ...
and the combination of bithional and
levamisole Levamisole, sold under the brand name Ergamisol among others, is a medication used to treat parasitic worm infections, specifically ascariasis and hookworm infections. It is taken by mouth. Side effects may include abdominal pain, vomiting, ...
. Most important commercial
anthelmintics Anthelmintics or antihelminthics are a group of antiparasitic drugs that expel parasitic worms ( helminths) and other internal parasites from the body by either stunning or killing them without causing significant damage to the host. They may als ...
are shown to be practically useless, including
albendazole Albendazole is a broad-spectrum antihelmintic and antiprotozoal agent of the benzimidazole type. It is used for the treatment of a variety of intestinal parasite infections, including ascariasis, pinworm infection, hookworm infection, trichuri ...
,
praziquantel Praziquantel, sold under the brandname Biltricide among others, is a medication used to treat a number of types of parasitic worm infections in mammals, birds, amphibians, reptiles, and fish. In humans specifically, it is used to treat schist ...
, nitroxynil,
triclabendazole Triclabendazole, sold under the brand name Egaten among others, is a medication used to treat fascioliasis and paragonimiasis. It is very effective for both conditions. Treatment in hospital may be required. It is taken by mouth with typicall ...
, profenophos and netobimin; while
niclosamide Niclosamide, sold under the brand name Niclocide among others, is an anthelmintic medication used to treat tapeworm infestations, including diphyllobothriasis, hymenolepiasis, and taeniasis. It is not effective against other worms such as ...
has high efficacy (99%) against immature fluke but not adult fluke, and 2-tertiary-butyl benzthiazole compound (CGA 72630),
hexachlorophene Hexachlorophene, also known as Nabac, is an organochlorine compound that was once widely used as a disinfectant. The compound occurs as a white odorless solid, although commercial samples can be off-white and possess a slightly phenolic odor. It ...
and resorantel are highly effective against both adult and immature flukes. An in vitro demonstration shows that