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''Paramphistomum cervi'', the
type species In 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 species that contains the biological type specimen ...
of ''
Paramphistomum ''Paramphistomum'' is a genus of parasitic flatworms belonging to the digenetic trematodes. It includes flukes which are mostly parasitising livestock ruminants, as well as some wild mammals. They are responsible for the serious disease called ...
'', is a
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
flat worm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegmen ...
belonging to the
class Class 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 differentl ...
Trematoda Trematoda is a Class (biology), class of flatworms known as flukes. They are obligate parasite, obligate internal Parasitism, parasites with a complex biological life cycle, life cycle requiring at least two Host_(biology), hosts. The intermedia ...
. It is a tiny
fluke Fluke may refer to: Biology * Fluke (fish), a species of marine flatfish * Fluke (tail), the lobes of the tail of a cetacean, such as dolphins or whales, ichthyosaurs, mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ...
mostly parasitising
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animals ...
ruminants Ruminants ( suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
, as well as some wild
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
. 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, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult mal ...
and
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals 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 domesticated s ...
. Its
symptoms Signs and symptoms are the observed or detectable signs, and experienced symptoms of an disease, illness, injury, or condition. A sign for example may be a higher or lower temperature than normal, raised or lowered blood pressure or an abnormali ...
include profuse
diarrhoea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin wi ...
,
anaemia Anemia or anaemia (British English) is a blood disorder in which the blood has a reduced ability to carry oxygen due to a lower than normal number of red blood cells, or a reduction in the amount of hemoglobin. When anemia comes on slowly, th ...
,
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, overwo ...
, 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 Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
,
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
,
Eastern Europe Eastern Europe is a subregion of the Europe, European continent. As a largely ambiguous term, it has a wide range of geopolitical, geographical, ethnic, cultural, and socio-economic connotations. The vast majority of the region is covered by Russ ...
, and
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
. It is most commonly found in
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
regions. The most debilitating cases are reported in Europe from
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedon ...
,
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
,
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
, and
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...
and also in Asia from
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
, and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. The parasitic infection was first described from
Punjab Punjab (; Punjabi: پنجاب ; ਪੰਜਾਬ ; ; also romanised as ''Panjāb'' or ''Panj-Āb'') is a geopolitical, cultural, and historical region in South Asia, specifically in the northern part of the Indian subcontinent, comprising ...
, India.


Description

Adult ''P. cervi'' is
conical A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
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 BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
. 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: Greece 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 ...
: ''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, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
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 als ...
s are present towards the posterior region of the body.
Testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testoster ...
are slightly lobed and are located anterior to the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
. Eggs are clear shell and measure about 140 x 80 µ; 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 birth to reproduction ending with the production of the offspring *Life-cycle hypothesis, ...
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 a ...
,
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...
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 a ...
, 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 alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
self-fertilises in the mammalian
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment allow ...
, and keep the eggs inside the
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
. 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 People *Jim Host (born 1937), American businessman * Michel Host ...
intestine and are expelled along with
faeces Feces ( or faeces), known colloquially and in slang as poo and poop, 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 large intestine. Feces contain a relati ...
. Eggs are deposited in water and hatch at optimal temperature of 27 °C in 12–17 days to become ciliated
miracidia Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two suckers: one ventral and the other oral. Trematodes are covered by a tegument, that protects the organism from the environment by providing secr ...
. 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 ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
''
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 ...
'', ''
Planorbis ''Planorbis'' is a genus of air-breathing freshwater snails, aquatic pulmonate gastropod mollusks in the family Planorbidae, the ram's horn snails, or planorbids. All species in this genus have sinistral or left-coiling shells.Bouchet, P.; Rosen ...
'', ''
Physa ''Physa'' is a genus of small, left-handed or sinistral, air-breathing freshwater snails, aquatic pulmonate gastropod mollusks in the subfamily Physinae of the family Physidae.MolluscaBase eds. (2021). MolluscaBase. Physa Draparnaud, 1801. Ac ...
'' '' Stagnicola'' 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 hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
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 µ). 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 Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two suckers: one ventral and the other oral. Trematodes are covered by a tegument, that protects the organism from the environment by providing secr ...
. 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, usually a bacterium or a protist or rarely an invertebrate animal, that helps the organism to survive in unfavorable environmental conditions. It can be thought of as a state of ...
to become
metacercariae Trematodes are parasitic flatworms of the class ''Trematoda'', specifically parasitic flukes with two suckers: one ventral and the other oral. Trematodes are covered by a tegument, that protects the organism from the environment by providing secr ...
. 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. ...
, 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 ingest ...
. Once they reach the
duodenum The duodenum is the first section of the small intestine in most higher vertebrates, including mammals, reptiles, and birds. In fish, the divisions of the small intestine are not as clear, and the terms anterior intestine or proximal intestine m ...
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 specialised for the absorption by enterocytes of small nutrient molecules which have been previous ...
, their
cyst A cyst is a closed sac, having a distinct envelope and cell division, division compared with the nearby Biological tissue, tissue. Hence, it is a cluster of Cell (biology), cells that have grouped together to form a sac (like the manner in which ...
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 GI tract contains all the major organs of the digestive system, in humans and ...
. 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 series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
. 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''), or gall, is a dark-green-to-yellowish-brown fluid produced by the liver of most vertebrates that aids the digestion of lipids in the small intestine. In humans, bile is produced continuously by the liver (liver bile ...
and
pancreatic The pancreas is an Organ (anatomy), organ of the digestive system and endocrine system of vertebrates. In humans, it is located in the abdominal cavity, 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 fro ...
s and non-pathogenic. However, they do cause the intestinal villi to erode and instil
inflammation Inflammation (from la, wikt:en:inflammatio#Latin, inflammatio) is part of the complex biological response of body tissues to harmful stimuli, such as pathogens, damaged cells, or Irritation, irritants, and is a protective response involving im ...
. 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, vagi ...
, discolouration,
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
, 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 organic tissue that results from cell proliferati ...
, and
fibrosis Fibrosis, also known as fibrotic scarring, is a pathological wound healing in which connective tissue replaces normal parenchymal tissue to the extent that it goes unchecked, leading to considerable tissue remodelling and the formation of perma ...
. Paramphistomiasis is responsible for severe economic losses to
milk Milk is a white liquid food produced by the mammary glands of mammals. It is the primary source of nutrition for young mammals (including breastfed human infants) before they are able to digestion, digest solid food. Immune factors and immune ...
,
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
and
wool Wool is the textile fibre 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 properties similar to animal wool. As ...
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, often referred to simply as anorexia, is an eating disorder characterized by low weight, food restriction, body image disturbance, fear of gaining weight, and an overpowering desire to be thin. ''Anorexia'' is a term of Gre ...
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. Alon ...
, clorsulon,
ivermectin Ivermectin (, '' EYE-vər-MEK-tin'') 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, today it is used to treat inf ...
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 and without causing significant damage to the host. They may a ...
are shown to be practically useless, including
albendazole Albendazole (also known as albendazolum) is a broad-spectrum anthelmintic and antiprotozoal agent of the benzimidazole type. It is used for the treatment of a variety of intestinal parasite infections, including ascariasis, pinworm infection, h ...
,
praziquantel Praziquantel (PZQ), sold under the brandname Biltricide among others, is a medication used to treat a number of types of helminthiasis, parasitic worm infections in mammals, birds, amphibians, reptiles, and fish. In humans specifically, it is us ...
, 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 typically ...
, 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 flukes ...
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