Eucestoda

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Eucestoda
Taenia saginata adult 5260 lores.jpg
Taenia saginata adult
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Platyhelminthes
Class: Cestoda
Subclass: Eucestoda

Eucestoda, commonly referred to as tapeworms, is the larger of the two subclasses of flatworms in the class Cestoda (the other subclass is Cestodaria). Larvae have six posterior hooks on the scolex (head), in contrast to the ten-hooked Cestodaria. All tapeworms are endoparasites of vertebrates, living in the digestive tract or related ducts. Examples are the pork tapeworm (Taenia solium) with a human definitive host, and pigs as the secondary host, and Moniezia expansa , the definitive hosts of which are ruminants.

Contents

Body structure

Adult Eucestoda have a white-opaque dorso-ventrally flattened appearance, and are elongated, ranging in length from a few millimeters (about ¼") to 25 meters (80'). [1] Almost all members, except members of the orders Caryophyllidea and Spathebothriidea, are polyzoic with repeated sets of reproductive organs down the body length, and almost all members, except members of the order Dioecocestidae, are protandral hermaphrodites. Most except caryophyllideans consist of a few to 4000 proglottids (segments) that show a characteristic body differentiation pattern into scolex (head), neck, and strobila. [2]

The scolex, located at the anterior end, is a small (usually less than 1 mm) holdfast organ with specific systems for fastening itself to materials: rostrum, acetabula, suckers, bothria, grooves, and hooks. The small neck region, directly behind the scolex, consists of an undifferentiated tissue region of proglottid proliferation, leading into a zone of increasing and continuous proglottid differentiation. As such, the main and largest section of the body, the strobila, consists of a chain of increasingly mature proglottids. These cytological processes are not well understood at present.

Members of the Eucestoda have no mouth or digestive tract, and instead absorb nutrients through a layer of microtriches over the tegument at the shared body wall surface. [3] In addition to the body wall, several other systems are common to the whole length of the tapeworm, including excretory canals, nerve fibers, and longitudinal muscles. [4] The excretory system is responsible for osmoregulation and consists of blind-ending flame bulbs communicating through a duct system. The nervous system, often referred to as a "ladder system," is a system of longitudinal connectives and transverse ring commissures. [5]

Reproduction

The reproductive systems develop progressively along the differentiated proglottids of the strobila region, with each proglottid developing one or two sets of sexual organs that differentiate at different times in a species-specific pattern, usually male-first. [5] Thus, moving in the posterior direction of the continuously maturing proglottid chain, there are proglottids with mature male reproductive organs, then proglottids with mature female reproductive organs, and then proglottids with fertilized eggs in the uterus, a condition commonly referred to as "gravid." [4]

Proglottids

An atrium on the lateral margin of each proglottid contains the openings to both the male and female genital ducts. Follicular testes produce sperm, which are carried by a system of ducts to the cirrus, an eversible copulatory organ that usually has a hypodermic system of spines and a holdfast system of hooks. The main specialized female reproductive organs are an ovary that produces eggs and a vitellarium that produces yolk cells. Yolk cells travel in a duct system to the oviduct, where, in a modified region, the ovum is enclosed in a shell with yolk cells. After the gonads and their ducts have finished maturing, the female reproductive organs begin to mature. The oviduct develops a vagina and enlarges into the uterus, where fertilization and embryonic development occur. [5]

Egg formation is a result of copulation. A proglottid can copulate with itself, with other proglottids in the same worm, or with proglottids in other worms, [2] and hypodermic fertilization sometimes occurs. [5] When a gravid proglottid that is distended with an embryo reaches the end of the strobila, it detaches and passes out of the host intact with feces, [4] with or without some tissue degeneration. [5] In the order Pseudophyllidea, the uterus has a pore and the proglottid sheds the shelled embryo, only becoming detached when exhausted. [5]

Some members of the Eucestoda (such as Echinococcus , Sparganum , Taenia multiceps sp., and Mesocestoides sp.) can reproduce asexually through budding, [5] which initiates a metagenesis of alternating sexually and asexually reproducing generations. [4]

Life stages

A tapeworm can live from a few days to over 20 years. [5] Eucestoda ontogenesis continues through metamorphosing in different larval stages inside different hosts. The initial six-hooked embryo, known as an oncosphere or hexacanth, forms through cleavage. In the order Pseudophyllidea, it remains enclosed in a ciliated embryophore. The embryo continues to develop in other host species, with two intermediate hosts generally needed. [5] It gains entry to its first intermediate host by being eaten.

Except for members of the order Taeniidae, the first intermediate host is an arthropod, and except for in the case of Archigetes spp. (which can attain sexual maturity in freshwater oligochaeta), the second host is usually a fish, but can be another invertebrate or vertebrate. [4] After the scolex has differentiated and matured in the larval stage, growth will stop until a vertebrate eats the intermediate host, and then the strobila develops. Adult tapeworms often have a high final host specificity, with some species only found in one host vertebrate. [5]

Common infective species

Final hostSpecies
Humans Diphyllobothrium latum , Spirometra erinacea euopaei , Taenia solium , T. saginata , T. asiatica , Mesocestoides leptothylacus (rarely), Rodentolepis (syn. Vampiro-, Hymenolepis ) nana , Hymenolepis diminuta
CatsDiphyllobothrium latum, Spirometra erinacea euopaei, T. pisiformis , Echinococcus multilocularis , Dipylidium caninum
DogsDiphyllobothrium latum, Spirometra erinacea euopaei, T. hydatigena , T. ovis , T. pisiformis , T. multiceps , T. serialis , E. granulosus , E. multilocaris , Mesocestoides leptothylacus , Dipylidium caninum
Fish Caryophyllaeus laticeps , Glaridacris catostomi , Archigetes sieboldi, Triaenophorus sp., Eubothrium sp., Rhinobothrium sp., Phyllobothrium sp., Proteocephalus ambloplites
Fish-Eating Birds Ligula intestinalis , Schistocephalus solidus
Rodents Schistocephalus solidus , Rodentolepis nana , Hymenolepis diminuta
Horses Anoplocephala magna , Anoplocephala perfoliata
Ruminants Moniezia expans , Avitellina sp., Thysaniezia sp., Stilesia sp.
Chickens Davainea proglottina , Raillietina tetragona , Amoebotaenia sp., Choanataenia sp., Hymenolepis carioca
Foxes T. ovis , T. multiceps , T. serialis , E. granulosus , E. multilocularis , Mesocestoides leptothylacus , Dipylidium caninum
Lagomorphs Cittotaenia sp. [4]

Medical importance

Taeniasis

Taeniasis is an infection within the intestines by adult tapeworms belonging to the genus Taenia . [6] [7] It is due to eating contaminated undercooked beef or pork. [6] There are generally no or only mild symptoms. [6] Symptoms may occasionally include weight loss or abdominal pain. [8] Segments of tapeworm may be seen in the stool. [8]

Cysticercosis

Cysticercosis is a tissue infection caused by the young form of the pork tapeworm. [9] [10] Infection occurs through swallowing or antiperistaltic contractions during regurgitation carrying eggs or gravid proglottids to the stomach. At this point, larvae hatch when exposed to enzymes and penetrate the intestinal wall, travelling through the body through blood vessels to tissues like the brain, the eye, muscles, and the nervous system (called neurocysticercosis). [5]

At these sites, the parasites lodge and form cysts, a condition called cysticercosis, producing inflammatory reactions and clinical issues when they die, sometimes causing serious or fatal damage. In the eye, the parasites can cause visual loss, and infection of the spine and adjacent leptomeninges can cause paresthesias, pain, or paralysis. [11]

Echinococcosis (hydatid disease)

Nomenclature framework for Cystic Echinococcosis surgery Vuitton et al - International consensus on terminology - parasite200043-fig3.png
Nomenclature framework for Cystic Echinococcosis surgery

Humans become accidental hosts to worms of the genus Echinococcus, playing no role in the worm's biological cycle. This can result in echinococcosis, also called hydatid disease. Humans (usually children) become infected by direct contact with dogs and eating food contaminated with dog feces. Common sites of infection are the liver, the lungs, muscles, bones, kidneys, and the spleen. [12]

Eggs hatch in the gastrointestinal tract after the consumption of contaminated food, after which the larvae travel to the liver through portal circulation. Here, the larvae are trapped and usually develop into hydatid cysts. While the liver is the first filter for trapping them, the lungs act as the second filter site, trapping most of the larvae that are not trapped by the liver. Some larvae escape from the lungs to cause cysts in other tissues. [5]

When a larva becomes established in tissue, it develops into a "bladderworm" or "hydatid" and can cause various cancer-like cysts that may rupture and interact with nearby organs. Most cases are asymptomatic, and the mortality rate is low, but various complications from these interactions may lead to debilitating illness.

Hymenolepiasis

Arthropods are intermediate hosts of Hymenolepis nana , otherwise known as the "dwarf tapeworm," while humans are used as final hosts. Humans become infected and develop hymenolepiasis through eating infected arthropods, ingesting eggs in water inhabited by arthropods, or from dirty hands. This is a common and widespread intestinal worm. [5]

While light infections are usually asymptomatic, autoinfection through eating the eggs of worms in the intestines is possible, and it can lead to hyperinfection. Humans can also become hyperinfected through ingesting grain products contaminated by infected insects. Infections involving more than two thousand worms can cause many different gastrointestinal symptoms and allergic responses. Common symptoms include chronic urticaria, skin eruption, and phlyctenular keratoconjunctivitis. [13]

Diphyllobothriasis

Diphyllobothriasis is caused by infection with Diphyllobothrium latum (also known as the "broad tapeworm" or "fish tapeworm") and related species. Humans become infected by eating raw, undercooked, or marinated fish acting as a second intermediate or paratenic host harboring metacestodes or plerocercoid larvae. [14]

Clinical symptoms are due to the large size of the tapeworm, which often reaches a length exceeding 15 m (49 ft). The most common symptom is pernicious anemia, caused by the absorption of vitamin B12 by the worm. [5] Other symptoms include various intestinal issues, slight leukocytosis, and eosinophilia.

Sparganosis

Sparganosis is caused by the plerocercoid larvae of the tapeworm Spirometra. Humans become infected by drinking contaminated water, eating raw or poorly cooked infected flesh, or from using poultices of raw infected flesh (usually raw pork or snake) on skin or mucous membranes. [15]

The most common symptom is a painful, slowly growing nodule in the subcutaneous tissues, which may migrate. Infection in the eye area can cause pain, irritation, edema, and excess watering. When the orbital tissues become infected, the swelling can cause blindness. An infected bowel may become perforated. Brain infection can cause granulomas, hematomas, and abscesses.

Subdivisions

The evolutionary history of the Eucestoda has been studied using ribosomal RNA, mitochondrial and other DNA, and morphological analysis and continues to be revised. "Tetraphyllidea" is seen to be paraphyletic; "Pseudophyllidea" has been broken up into two orders, Bothriocephalidea and Diphyllobothriidea. [16] [17] [18] Hosts, whose phylogeny often mirrors that of the parasites (Fahrenholz's rule), are indicated in italics and parentheses, the life-cycle sequence (where known) shown by arrows as (intermediate host1 [→ intermediate host2] → definitive host). Alternatives, generally for different species within an order, are shown in square brackets.

Cestoda

Gyrocotylidea (fishes)

Amphilinidea (crustaceans → fishes/turtles)

Eucestoda

Spathebothriidea (amphipods → fishes)

Caryophyllidea (annelids → fishes)

Haplobothriidea (freshwater fishes → bowfin)

Diphyllobothriidea (copepods [→ fishes] → mammals)

Diphyllidea (elasmobranchs inc. rays, sharks)

Trypanorhyncha (fishes/crustaceans/molluscs → bony fishes/selachians)

Bothriocephalidea (crustaceans [→ teleost] → teleost fishes/amphibians)

Litobothriidea (lamniform sharks)

Lecanicephalidea (molluscs → selachians)

Rhinebothriidea (stingrays)

"Tetraphyllidea" (copepods → fishes/decapods/cephalopods → selachians)

"Tetraphyllidea"

Onchoproteocephalidea (crustaceans → inverts/verts → fishes/amphibians/reptiles)

Nippotaeniidea (crustaceans → fishes)

Mesocestoididae (mammals/birds)

Tetrabothriidea (crustaceans?/cephalopods?/teleosts? → seabirds/cetaceans/pinnipeds)

Cyclophyllidea (mammals → mammals, or insects → birds)

tapeworms

Related Research Articles

<i>Diphyllobothrium</i> Genus of flatworms

Diphyllobothrium is a genus of tapeworms which can cause diphyllobothriasis in humans through consumption of raw or undercooked fish. The principal species causing diphyllobothriasis is D. latum, known as the broad or fish tapeworm, or broad fish tapeworm. D. latum is a pseudophyllid cestode that infects fish and mammals. D. latum is native to Scandinavia, western Russia, and the Baltics, though it is now also present in North America, especially the Pacific Northwest. In Far East Russia, D. klebanovskii, having Pacific salmon as its second intermediate host, was identified.

<span class="mw-page-title-main">Hymenolepiasis</span> Medical condition

Hymenolepiasis is infestation by one of two species of tapeworm: Hymenolepis nana or H. diminuta. Alternative names are dwarf tapeworm infection and rat tapeworm infection. The disease is a type of helminthiasis which is classified as a neglected tropical disease.

<i>Taenia solium</i> Species of Cestoda

Taenia solium, the pork tapeworm, belongs to the cyclophyllid cestode family Taeniidae. It is found throughout the world and is most common in countries where pork is eaten. It is a tapeworm that uses humans as its definitive host and pigs as the intermediate or secondary hosts. It is transmitted to pigs through human feces that contain the parasite eggs and contaminate their fodder. Pigs ingest the eggs, which develop into larvae, then into oncospheres, and ultimately into infective tapeworm cysts, called cysticercus. Humans acquire the cysts through consumption of uncooked or under-cooked pork and the cysts grow into adult worms in the small intestine.

<i>Taenia</i> (flatworm) Genus of flatworms

Taenia is the type genus of the Taeniidae family of tapeworms. It includes some important parasites of livestock. Members of the genus are responsible for taeniasis and cysticercosis in humans, which are types of helminthiasis belonging to the group of neglected tropical diseases. More than 100 species are recorded. They are morphologically characterized by a ribbon-like body composed of a series of segments called proglottids; hence the name Taenia. The anterior end of the body is the scolex. Some members of the genus Taenia have an armed scolex ; of the two major human parasites, Taenia saginata has an unarmed scolex, while Taenia solium has an armed scolex.

<i>Taenia saginata</i> Species of flatworm

Taenia saginata, commonly known as the beef tapeworm, is a zoonotic tapeworm belonging to the order Cyclophyllidea and genus Taenia. It is an intestinal parasite in humans causing taeniasis and cysticercosis in cattle. Cattle are the intermediate hosts, where larval development occurs, while humans are definitive hosts harbouring the adult worms. It is found globally and most prevalently where cattle are raised and beef is consumed. It is relatively common in Africa, Europe, Southeast Asia, South Asia, and Latin America. Humans are generally infected as a result of eating raw or undercooked beef which contains the infective larvae, called cysticerci. As hermaphrodites, each body segment called proglottid has complete sets of both male and female reproductive systems. Thus, reproduction is by self-fertilisation. From humans, embryonated eggs, called oncospheres, are released with faeces and are transmitted to cattle through contaminated fodder. Oncospheres develop inside muscle, liver, and lungs of cattle into infective cysticerci.

<i>Echinococcus granulosus</i> Species of flatworm

Echinococcus granulosus, also called the hydatid worm or dog tapeworm, is a cyclophyllid cestode that dwells in the small intestine of canids as an adult, but which has important intermediate hosts such as livestock and humans, where it causes cystic echinococcosis, also known as hydatid disease. The adult tapeworm ranges in length from 3 mm to 6 mm and has three proglottids ("segments") when intact—an immature proglottid, mature proglottid and a gravid proglottid. The average number of eggs per gravid proglottid is 823. Like all cyclophyllideans, E. granulosus has four suckers on its scolex ("head"), and E. granulosus also has a rostellum with hooks. Several strains of E. granulosus have been identified, and all but two are noted to be infective in humans.

<i>Dipylidium caninum</i> Species of flatworm

Dipylidium caninum, also called the flea tapeworm, double-pored tapeworm, or cucumber tapeworm is a cyclophyllid cestode that infects organisms afflicted with fleas and canine chewing lice, including dogs, cats, and sometimes human pet-owners, especially children.

<i>Taenia pisiformis</i> Species of flatworm

Taenia pisiformis, commonly called the rabbit tapeworm, is an endoparasitic tapeworm which causes infection in lagomorphs, rodents, and carnivores. Adult T. pisiformis typically occur within the small intestines of the definitive hosts, the carnivores. Lagomorphs, the intermediate hosts, are infected by fecal contamination of grasses and other food sources by the definitive hosts. The larval stage is often referred to as Cysticercus pisiformis and is found on the livers and peritoneal cavities of the intermediate hosts. T. pisiformis can be found worldwide.

<span class="mw-page-title-main">Taeniasis</span> Parasitic disease due to infection with tapeworms belonging to the genus Taenia

Taeniasis is an infection within the intestines by adult tapeworms belonging to the genus Taenia. There are generally no or only mild symptoms. Symptoms may occasionally include weight loss or abdominal pain. Segments of tapeworm may be seen in the stool. Complications of pork tapeworm may include cysticercosis.

Spirometra erinaceieuropaei is a parasitic tapeworm that infects domestic animals and humans. The medical term for this infection in humans and other animals is sparganosis. Morphologically, these worms are similar to other worms in the genus Spirometra. They have a long body consisting of three sections: the scolex, the neck, and the strobilia. They have a complex life cycle that consists of three hosts, and can live in varying environments and bodily tissues. Humans can contract this parasite in three main ways. Historically, humans are considered a paratenic host; however, the first case of an adult S. erinaceieuropaei infection in humans was reported in 2017. Spirometra tapeworms exist worldwide and infection is common in animals, but S. erinaceieuropaei infections are rare in humans. Treatment for infection typically includes surgical removal and anti-worm medication.

<i>Hymenolepis</i> (flatworm) Genus of worms

Hymenolepis is a genus of cyclophyllid tapeworms that cause hymenolepiasis. They parasitise mammals, including humans. Some notable species are:

<span class="mw-page-title-main">Cestoda</span> Class of flatworms

Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, known as tapeworms. Their bodies consist of many similar units known as proglottids—essentially packages of eggs which are regularly shed into the environment to infect other organisms. Species of the other subclass, Cestodaria, are mainly fish infecting parasites.

Diphyllobothrium mansonoides is a species of tapeworm (cestodes) that is endemic to North America. Infection with D. mansonoides in humans can result in sparganosis. Justus F. Mueller first reported this organism in 1935. D. mansonoides is similar to D. latum and Spirometra erinacei. When the organism was discovered, scientist did not know if D. mansonoides and S. erinacei were separate species. PCR analysis of the two worms has shown the two to be separate but closely related organisms.

<i>Raillietina</i> Genus of flatworms

Raillietina is a genus of tapeworms that includes helminth parasites of vertebrates, mostly of birds. The genus was named in 1920 in honour of a French veterinarian and helminthologist, Louis-Joseph Alcide Railliet. Of the 37 species recorded under the genus, Raillietina demerariensis, R. asiatica, and R. formsana are the only species reported from humans, while the rest are found in birds. R. echinobothrida, R. tetragona, and R. cesticillus are the most important species in terms of prevalence and pathogenicity among wild and domestic birds.

<i>Raillietina echinobothrida</i> Species of flatworm

Raillietina echinobothrida is a parasitic tapeworm belonging to the class Cestoda. It is the most prevalent and pathogenic helminth parasite in birds, particularly in domestic fowl, Gallus domesticus Linnaeus, 1758. It requires two hosts, birds and ants, for completion of its life cycle. It is a hermaphrodite worm having both the male and female reproductive organs in its body. The parasite is responsible for 'nodular tapeworm disease' in poultry.

Taenia asiatica, commonly known as Asian taenia or Asian tapeworm, is a parasitic tapeworm of humans and pigs. It is one of the three species of Taenia infecting humans and causes taeniasis. Discovered only in 1980s from Taiwan and other East Asian countries as an unusual species, it is so notoriously similar to Taenia saginata, the beef tapeworm, that it was for a time regarded as a slightly different strain. But anomaly arose as the tapeworm is not of cattle origin, but of pigs. Morphological details also showed significant variations, such as presence of rostellar hooks, shorter body, and fewer body segments. The scientific name designated was then Asian T. saginata. But the taxonomic consensus turns out to be that it is a unique species. It was in 1993 that two Korean parasitologists, Keeseon S. Eom and Han Jong Rim, provided the biological bases for classifying it into a separate species. The use of mitochondrial genome sequence and molecular phylogeny in the late 2000s established the taxonomic status.

<i>Raillietina tetragona</i> Species of flatworm

Raillietina tetragona is a parasitic tapeworm belonging to the class Cestoda. It is a cosmopolitan helminth of the small intestine of pigeon, chicken and guinea fowl, and is found throughout the world.

<i>Taenia hydatigena</i> Species of flatworm

Taenia hydatigena is one of the adult forms of the canine and feline tapeworm. This infection has a worldwide geographic distribution. Humans with taeniasis can infect other humans or animal intermediate hosts by eggs and gravid proglottids passed in the feces.

<span class="mw-page-title-main">Cysticercus</span> Larval tapeworm

Cysticercus is a scientific name given to the young tapeworms (larvae) belonging to the genus Taenia. It is a small, sac-like vesicle resembling a bladder; hence, it is also known as bladder worm. It is filled with fluid, in which the main body of the larva, called scolex, resides. It normally develops from the eggs, which are ingested by the intermediate hosts, such as pigs and cattle. The tissue infection is called cysticercosis. Inside such hosts, they settle in the muscles. When humans eat raw or undercooked pork or beef that is contaminated with cysticerci, the larvae grow into adult worms inside the intestine. Under certain circumstances, specifically for the pork tapeworm, the eggs can be accidentally eaten by humans through contaminated foodstuffs. In such case, the eggs hatch inside the body, generally moving to muscles as well as inside the brain. Such brain infection can lead to a serious medical condition called neurocysticercosis. This disease is the leading cause of acquired epilepsy.

<i>Echinococcus vogeli</i> Species of flatworm

Echinococcus vogeli is a small cyclophyllid tapeworm found in Central and South America. E. vogeli, as well as other members of the genus Echinococcus, produce a disease called echinococcosis. Echinococcosis, also known has hydatidosis, is a result of ingesting the eggs of the genus Echinococcus. E. vogeli is similar to E. multilocularis in that both species produces many small cysts that spread throughout the internal organs of the infected animal. The ingestion of E. vogeli eggs, and the spreading of the cysts through infected host, will results in Polycystic Echinococcosis.

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