View source for Acanthocephala

{{otheruses}}
{{Taxobox
| color = pink
| name = Acanthocephala
| image = C wegeneri.JPG
| image_width = 250px
| image_caption = ''Corynosoma wegeneri''
| regnum = [[Animal]]ia
| subregnum = [[Eumetazoa]]
| unranked_phylum = [[Bilateria]]
| superphylum = [[Platyzoa]]
| phylum = '''Acanthocephala'''
| phylum_authority = Kohlreuther, 1771
| subdivision_ranks = [[Class (biology)|Classes]]
| subdivision = 
[[Archiacanthocephala]]<br/>
[[Palaeacanthocephala]]<br/>
[[Eoacanthocephala]]
}}
The '''Acanthocephala''' (gr. ''Acanthus'' — thorn ''Kephale'' — head) is a [[phylum (biology)|phylum]] of [[parasitic]] worms, characterised by the presence of an evertable [[proboscis]], armed with spines, which it uses to pierce and hold the [[gut]] wall of its host. Acanthocephalans typically have [[Parasitic life cycles|complex life cycles]], involving a number of hosts, including [[invertebrate]]s, [[fish]]es, [[amphibia]]ns, [[bird]]s, and [[mammal]]s. About 1150 [[species]] have been described.

==Morphological characteristics==

There are several morphological characteristics that distinguish acanthocephalans from other phyla of parasitic worms.

===Digestion===
Acanthocephalans lack a [[mouth]] or [[alimentary canal]]. This is a feature they share with the [[cestoda]] (tapeworms), although the two groups are not related. Adult stages live in the [[intestine]]s of their host and uptake nutrients which have been [[digest]]ed by the host, directly, through their body surface.

===Proboscis===
The most notable feature of the acanthocephala is the presence of an [[anterior]], protrudible [[proboscis]] that is usually covered with spiny hooks (hence the common name: thorny headed worm). The proboscis bears rings of recurved hooks arranged in horizontal rows, and it is by means of these hooks that the animal attaches itself to the tissues of its host. The hooks may be of two or three shapes, usually, longer, more slender hooks are arranged along the length of the proboscis, with several rows of more sturdy, shorter nasal hooks around the base of the proboscis. The proboscis is used to pierce the gut wall of the final host, and hold the parasite fast while it completes its life cycle.
Like the body, the proboscis is hollow, and its cavity is separated from the body cavity by a ''septum'' or ''proboscis sheath''. Traversing the cavity of the proboscis are [[muscle]]-strands inserted into the tip of the proboscis at one end and into the septum at the other.  Their contraction causes the proboscis to be invaginated into its cavity. The whole proboscis apparatus can also be, at least partially, withdrawn into the body cavity, and this is effected by two retractor muscles which run from the posterior aspect of the septum to the body wall.

[[Image:Corynosoma drawing.jpg|thumb|left|Diagram of Acanthocephalan morphology|Some key features of acanthocephalan morphology]]

===Phylogenetic relationships===
Acanthocephalans are highly adapted to a parasitic mode of life, and have lost many organs and structures through evolutionary processes. This makes determining relationships with other higher taxa through morphological comparison problematic. [[Phylogeny|Phylogenetic]] analysis of the 18S [[ribosome|ribosomal]] gene has revealed that the Acanthocephala are most closely related to the [[rotifer]]s, or may even belong in that phylum.  The two are included among the [[Platyzoa]].

===Size===
The size of the animals varies greatly, from forms a few [[millimetre]]s in length to ''Gigantorhynchus gigas'', which measures from 100 to 650 mm.

===Skin===
The body surface of the acanthocephala is peculiar.  Externally, the skin has a thin [[cuticle]] covering  the [[Squamous epithelium|epidermis]], which consists of a [[syncytium]] with no [[cell wall]]s.  The syncytium is traversed by a series of branching [[tubule]]s containing fluid and is controlled by a few wandering, [[amoeboid]] [[cell nucleus|nuclei]]. Inside the syncytium is an irregular layer of circular muscle fibres, and within this again some rather scattered longitudinal fibres; there is no [[endothelium]].  In their micro-structure the muscular fibres resemble those of [[nematode]]s.

Except for the absence of the longitudinal fibres the skin of the proboscis resembles that of the body, but the fluid-containing tubules of the proboscis are shut off from those of the body. The canals of the proboscis open into a circular vessel which runs round its base. From the circular canal two sac-like projections called the ''lemnisci'' run into the cavity of the body, alongside the proboscis cavity. Each consists of a prolongation of the syncytial material of the proboscis skin, penetrated by canals and sheathed with a muscular coat. They seem to act as reservoirs into which the fluid which is used to keep the proboscis "erect" can withdraw when it is retracted, and from which the fluid can be driven out when it is wished to expand the proboscis.

===Nervous system===

The central ganglion of the nervous system lies behind the proboscis sheath or septum. It innervates the proboscis and projects two stout trunks posteriorly which supply the body.  Each of these trunks is surrounded by muscles, and this nerve-muscle complex is called a ''retinaculum''. In the male at least there is also a [[genital]] [[ganglion]]. Some scattered [[papilla]]e may possibly be sense-organs.

==="Brain-jacking"===

Thorny-headed worms begin their life cycle inside invertebrates that reside in lakes and rivers. ''Gammarus lacustris'', a small crustacean that feeds near ponds and rivers, is one invertebrate that the thorny-headed worm may occupy. This crustacean is predated by ducks and hides by avoiding light and staying away from the surface. However, when infected by a thorny-headed worm it becomes attracted toward light and surfaces itself. ''Gammarus lacustris'' will even go so far as to find a rock or a plant on the surface, clamp its mouth down, and latch on, making it easy prey for the duck.

It is thought that when ''Gammarus lacustris'' is infected with a thorny-headed worm, the parasite causes serotonin to be massively expressed. Serotonin is a neurotransmitter involved in emotions and mood. Researchers have found that during mating ''Gammarus lacustris'' expresses high levels of serotonin. Also during mating, the male ''Gammarus lacustris'' clamps down on the female and holds on for days. Researchers have additionally found that blocking serotonin releases clamping. Another experiment found that serotonin also reduces the photophobic behavior in ''Gammarus lacustris''. Thus, it is thought that the thorny-headed worm physiologically changes the behavior of the ''Gammarus lacustris'' in order to enter its final host, the bird.

===Sex===

The Acanthocephala are [[dioecious]]. There is a structure called the ''genital ligament'' which runs from the posterior end of the proboscis sheath to the posterior end of the body. In the male, two [[testes]] lie on either side of this. Each opens in a [[vas deferens]] which bears three [[diverticula]] or ''vesiculae seminales''. The male also possesses three pairs of cement glands, found behind the testes, which pour their secretions through a duct into the vasa deferentia. These unite and end in a [[penis]] which opens posteriorly.

In the female, the [[ovaries]] are found, like the testes, as rounded bodies along the ligament.  From these masses of [[ova]] dehisce into the body cavity and float in its fluid. Here the eggs are [[fertilize]]d and segment so that the young [[embryo]]s are formed within their mother's body.  The embryos escape into the [[uterus]] through the ''uterine bell'', a funnel like opening continuous with the uterus.  At the junction of the bell and the uterus there is a second small opening situated [[Dorsum (biology)|dorsal]]ly. The bell "swallows" the matured embryos and passes them on into the uterus, and from there, out of the body via the [[oviduct]]. Should the bell swallow any of the ova, or even one of the younger embryos, these are passed back into the body cavity through the second, dorsal, opening.

The embryo passes from the body of the female into the alimentary canal of the host and leaves this with the [[feces]].

===Other features===

A curious feature shared by both larva and adult is the large size of many of the cells, e.g. the nerve cells and cells forming the uterine bell. [[Polyploid]]y is common, with up to 343n having been recorded in some species. The acanthocephalans lack an excretory system, although some species have been shown to possess [[flame cell]]s (protonephridia).

==History==

The earliest recognisable description of Acanthocephala - a worm with a proboscis armed with hooks - was made by Italian author [[Francesco Redi]] (1684). In [[1771]] [[Koelreuther]] proposed the name Acanthocephala. Muller independetly called them Echinorhynchus in [[1776]]. [[Karl Rudolphi|Rudolphi]] in [[1809]] formally named them Acanthocephala.

Currently the phylum is divided into four classes - Palaeacanthocephala, Archiacanthocephala, Polyacanthocephala and Eoacanthocephala.

==Life cycles==
===General patterns===
Acanthocephalans have complex life cycles, involving a number of hosts, for both [[morphogenesis|development]]al and resting stages. Complete life cycles have been worked out for only 25 species.
Having been expelled by the female, the acanthocephalan embryo is released along with the feces of the host. For development to occur, the embryo needs to be ingested by an invertebrate, almost always a [[crustacean]] (there is one known life cycle which uses a [[mollusc]] as a first intermediate host). Inside the intermediate host, the acanthocephalan penetrates the gut wall, moves into the body cavity, encysts, and begins transformation into the infective ''cystacanth'' stage. This form has all the organs of the adult save the reproductive ones.
The parasite is released when the first intermediate host is ingested. This can be by a suitable final host, in which case the cystacanth develops into a mature adult, or by a [[paratenic]] host, in which the parasite again forms a cyst. When consumed by a suitable final host, the cycstacant ''excysts'', everts its proboscis and pierces the gut wall. It then feeds, grows and develops its sexual organs. Adult worms then mate. The male uses the excretions of its ''cement glands'' to plug the [[vagina]] of the female, preventing subsequent matings from occurring. Embryos develop inside the female, and the life cycle repeats.


===An example - ''Polymorphus'' spp.===
''Polymorphus'' spp. are parasites of [[seabirds]], particularly the [[Eider Duck]] (''Somateria mollissima''). Heavy infections of up to 750 parasites per bird are common, causing [[ulcer]]ation to the gut, disease and seasonal mortality. Recent research has suggested that there is no evidence of [[pathogen]]icity of ''Polymorphus'' spp. to intermediate crab hosts. The cystacanth stage is long lived and probably remains infective throughout the life of the crab.

The life cycle of ''Polymorphus'' spp. normally occurs between sea ducks (e.g. eiders and [[scoter]]s) and small crabs. Infections found in commercial-sized [[lobster]]s in Canada were probably acquired from crabs that form an important dietary item of lobsters. Cystacanths occurring in lobsters can cause economic loss to fishermen. There are no known methods of prevention or control.

==See also==
{{wikispecies|Acanthocephala}}
* [[Cestoda]]
* [[Digenea]]
* [[Monogenea]]

==References==
*Lühe, M. (1904). 'Geschichte und Ergebnisse der Echinorhynchen-Forschung bis auf Westrumb (1821)', Zoologische Ann.... Zeitschift, 1, 139-250.
*Amin, O. M. (1987). Key to the families and subfamilies of Acanthocephala, with erection of a new class (Polyacanthocephala) and a new order (Polyacanthorhynchida). Journal of Parasitology, 73, 1216-1219.
*Zimmer, C. 'Parasite Rex: Inside the Bizarre World of Nature's Most Dangerous Creatures' 92. IBSN-13: 978-0-7432-0011-0 ISBN-10: 0-7432-0011-X.
*Helluy & Thomas 2002 Proc. Royal.
*Tain, L, et al. "Altered Host Behavior and Brain Serotonergic Activity Caused by Acanthocephalans; Evidence for Specificity." The Royal Society, 2006.

==External links==
{{Wikisource1911Enc|Acamthocephala|Acanthocephala}}
* [http://www.fishdisease.net Fishdisease.net]
* [http://parasitology.informatik.uni-wuerzburg.de/login/n/h/0008.html Review]
* [http://www.rtis.com/nat/user/elsberry/taxa/acanth.htm Review]
* [http://ryoko.biosci.ohio-state.edu/~parasite/taxonomic_aaa.html Images]

[[Category:Acanthocephalans| ]]
[[Category:Veterinary parasitology]]
[[Category:Parasites]]

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