Cestoda

All 6,000 species of Cestoda are parasites, mainly intestinal; their definitive hosts are vertebrates, both terrestrial and marine, while their intermediate hosts include insects, crustaceans, molluscs, and annelids as well as other vertebrates. T. saginata, the beef tapeworm, can grow up to 20 m (65 ft); the largest species, the whale tapeworm Tetragonoporus calyptocephalus, can grow to over 30 m (100 ft). Species with small hosts tend to be small. For example, vole and lemming tapeworms are only in length, and those parasitizing shrews only . ==Anatomy==

Cestodes have no gut or mouth through which gas exchange also takes place. and allows it to transfer molecules back to the host. The Cestodaria have 10 larval hooks while Eucestoda have 6 larval hooks. Scolex '' with hooks and acetabula (suckers) present The scolex, which attaches to the intestine of the definitive host, is often minute in comparison with the proglottids. It is typically a four-sided knob, armed with suckers or hooks or both. Other species have ruffled or leaflike scolices, and there may be other structures to aid attachment. Body systems Circular and longitudinal muscles lie under the neodermis, beneath which further longitudinal, dorso-ventral and transverse muscles surround the central parenchyma. Protonephridial cells drain into the parenchyma. There are four longitudinal collection canals, two dorso-lateral and two ventro-lateral, running along the length of the worm, with a transverse canal linking the ventral ones at the posterior of each segment. When the proglottids begin to detach, these canals open to the exterior through the terminal segment. Cestodes are unable to synthesise lipids, which they use for reproduction, and are therefore entirely dependent on their hosts. The tapeworm body is composed of a series of segments called proglottids. These are produced from the neck by mitotic growth, which is followed by transverse constriction. The segments become larger and more mature as they are displaced backwards by newer segments. Mature proglottids are essentially bags of eggs, each of which is infective to the proper intermediate host. They are released and leave the host in feces, or migrate outwards as independent motile proglottids. ==Reproduction==

Cestodes are exclusively hermaphrodites, with both male and female reproductive systems in each body. The reproductive system includes one or more testes, cirri, vas deferens, and seminal vesicles as male organs, and a single lobed or unlobed ovary with the connecting oviduct and uterus as female organs. The common external opening for both male and female reproductive systems is known as the genital pore, which is situated at the surface opening of the cup-shaped atrium. Though they are sexually hermaphroditic and cross-fertilization is the norm, self-fertilization sometimes occurs and makes possible the reproduction of a worm when it is the only individual in its host's gut. During copulation, the cirri of one individual connect with those of the other through the genital pore, and then spermatozoa are exchanged. ==Life cycle==

'': Inset 5 shows the scolex, a disk with hooks on the end. Inset 6 shows the tapeworm's whole body, in which the scolex is the tiny, round tip in the top left corner, and a mature proglottid has just detached. '' relies on at least three hosts, crustaceans, fish, and humans. Other fish-eating mammals like bears can equally serve as definitive hosts. Cestodes are parasites of vertebrates, with each species infecting a single definitive host or group of closely related host species. All but amphilinids and gyrocotylids (which burrow through the gut or body wall to reach the coelom No asexual phases occur in the life cycle, as they do in other flatworms, but the life cycle pattern has been a crucial criterion for assessing evolution among Platyhelminthes. Cestodes produce large numbers of eggs, but each one has a low probability of finding a host. To increase their chances, different species have adopted various strategies of egg release. In the Pseudophyllidea, many eggs are released in the brief period when their aquatic intermediate hosts are abundant (semelparity). In contrast, in the terrestrial Cyclophyllidea, proglottids are released steadily over a period of years, or as long as their host lives (iteroparity). Another strategy is to have very long-lived larvae; for example, in Echinococcus, the hydatid larvae can survive for ten years or more in humans and other vertebrate hosts, giving the tapeworm an exceptionally long time window in which to find another host. Many tapeworms have a two-phase life cycle with two types of host. The adult Taenia saginata lives in the gut of a primate such as a human, its definitive host. Proglottids leave the body through the anus and fall to the ground, where they may be eaten with grass by a grazing animal such as a cow. This animal then becomes an intermediate host, the oncosphere boring through the gut wall and migrating to another part of the body such as the muscle. Here it encysts, forming a cysticercus. The parasite completes its life cycle when the intermediate host passes on the parasite to the definitive host, usually when the definitive host eats contaminated parts of the intermediate host, for example a human eating raw or undercooked meat. Diphyllobothrium exhibits a more complex, three-phase life cycle. If the eggs are laid in water, they develop into free-swimming oncosphere larvae. After ingestion by a suitable freshwater crustacean such as a copepod, the first intermediate host, they develop into procercoid larvae. When the copepod is eaten by a suitable second intermediate host, typically a minnow or other small freshwater fish, the procercoid larvae migrate into the fish's flesh where they develop into plerocercoid larvae. These are the infective stages for the mammalian definitive host. If the small fish is eaten by a predatory fish, its muscles too can become infected. ==Host immunity==

Hosts can become immune to infection by a cestode if the lining, the mucosa, of the gut is damaged. This exposes the host's immune system to cestode antigens, enabling the host to mount an antibody defence. Host antibodies can kill or limit cestode infection by damaging their digestive enzymes, which reduces their ability to feed and therefore to grow and to reproduce; by binding to their bodies; and by neutralising toxins that they produce. When cestodes feed passively in the gut, they do not provoke an antibody reaction. ==Evolution and phylogeny==

Fossil history Parasite fossils are rare, but recognizable clusters of cestode eggs, some with an operculum (lid) indicating that they had not erupted, one with a developing larva, have been discovered in fossil shark coprolites dating to the Permian, some 270 million years ago. The fossil organism Rugosusivitta, which was found in China at base of the Cambrian deposits in Yunnan just above the Ediacaran-Cambrian border, has great similarities to present day Cestodians. If correct, this would be the earliest example of a Platyzoan and also one of the earliest bilaterian body-fossils and might thus provide an insight to the living mode of Cestodians before they became specialized parasites. External The position of the Cestoda within the Platyhelminthes and other Spiralian phyla based on genomic analysis is shown in the phylogenetic tree. The non-parasitic flatworms, traditionally grouped as the "Turbellaria", are paraphyletic, as the parasitic Neodermata including the Cestoda arose within that grouping. The approximate times when major groups first appeared is shown in millions of years ago. }} Internal : body flatwormlike, not divided into proglottids : body wormlike, not divided into proglottids ": elaborate four-leaved scolex The evolutionary history of the Cestoda 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. 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. ==Interactions with humans==

drugs, some effective against many kinds of parasite, others more specific; these can be used both preventatively and to treat infections. For example, praziquantel is an effective treatment for tapeworm infection, and is preferred over the older niclosamide. While accidental tapeworm infections in developed countries are quite rare, such infections are more likely to occur in countries with poor sanitation facilities or where food hygiene standards are low. In Medieval times, in The Canon of Medicine, completed in 1025, the Persian physician Avicenna recorded parasites including tapeworms. Tapeworms are prominently mentioned in the System of a Down song "Needles": their inclusion within the song resulted in a lyrical dispute among band members. There are unproven claims that, around 1900, tapeworm eggs were marketed to the public as slimming tablets. A full-page coloured image, purportedly from a women's magazine of that period, reads "Fat: the enemy ... that is banished! How? With sanitized tape worms. Jar packed. No ill effects!" When television presenter Michael Mosley deliberately infected himself with tapeworms he gained weight due to increased appetite. Dieters still sometimes risk intentional infection, evidenced by a 2013 warning on American television. ==Notes==