Trematoda

Trematodes are commonly referred to as flukes. This term can be traced back to the Old English name for flounder, and refers to the flattened, rhomboidal shape of the organisms.

There are 18,000[1] to 24,000[2] known species of trematodes, divided into two subclasses — the Aspidogastrea and the Digenea. Aspidogastrea is the smaller subclass, comprising 61 species. These flukes mainly infect bivalves and bony fishes.[3] Digenea — which comprise the majority of trematodes — are found in certain mollusks and vertebrates.

Flukes that cause disease in humans are often classified based on the organ system they infect. For example:

• Blood flukes inhabit the blood in some stages of their life cycle. Blood flukes that cause disease in humans include Trichobilharzia regenti, which causes swimmer's itch, and seven species of genus Schistosoma which cause schistosomiasis: S. guineensis, S.haematobium, S. intercalatum, S. japonicum, S. malayensis, S. mansoni, S. mekongi. As a definitive host, humans are infected when the cercariae (the larval forms of trematodes) penetrate the skin. Any contact with water containing these cercariae can potentially result in infection. Adult blood flukes can live for years in human or animal reservoir hosts.[4] S.haematobium and S. japonicum are of particular importance, as these are carcinogenic parasites. S.haematobium, which infects the urinary bladder, is among the most important causes of bladder cancer in humans.[5][6] This organism is classified by the International Agency for Research on Cancer (IARC) as a Group 1 (extensively proven) carcinogen.[7] S. japonicum is associated with the development of liver cancer, and is classified as a Group 2B (possibly carcinogenic to humans) carcinogen.[7]

• Liver flukes are commonly found within bile ducts, liver, and gallbladder in certain mammalian and avian species. They include Clonorchis sinensis, Dicrocoelium dendriticum, Dicrocoelium hospes, Fasciola gigantica, Fasciola hepatica, Opisthorchis felineus, and Opisthorchis viverrini. Clonorchis and Opisthorchis are carcinogenic parasites that are strongly associated with the development of cancer of the bile ducts.[8][9]

• Lung flukes: there are ten species of lung flukes that infect humans, causing paragonimiasis.[10] Of these, the most common cause of human paragonimiasis is Paragonimus westermani, the oriental lung fluke.[11] Lung flukes require three different hosts in order to complete their life cycle. The first intermediate host is a snail, the second intermediate host is a crab or crayfish, and the definitive host for lung flukes is an animal or human host.[4]

• Intestinal flukes inhabit the epithelium of the small intestine. These include Fasciolopsis buski (which causes fasciolopsiasis), Metagonimus miyatai, Metagonimus takahashii, Metagonimus yokogawai (which cause metagonimiasis), and Heterophyes heterophyes and Heterophyes nocens (which cause heterophyiasis).

Varied trematodes, from 1911 Encyclopædia Britannica

Trematodes are flattened oval or worm-like animals, usually no more than a few centimeters in length, although species as small as 1 millimetre (0.039 in) are known. Their most distinctive external feature is the presence of two suckers, one close to the mouth, and the other on the underside of the animal.[12]

The body surface of trematodes comprises a tough syncytial tegument, which helps protect against digestive enzymes in those species that inhabit the gut of larger animals. It is also the surface of gas exchange; there are no respiratory organs.[12]

The mouth is located at the forward end of the animal, and opens into a muscular, pumping pharynx. The pharynx connects, via a short oesophagus, to one or two blind-ending caeca, which occupy most of the length of the body. In some species, the caeca are themselves branched. As in other flatworms, there is no anus, and waste material must be egested through the mouth.[12]

Although the excretion of nitrogenous waste occurs mostly through the tegument, trematodes do possess an excretory system, which is instead mainly concerned with osmoregulation. This consists of two or more protonephridia, with those on each side of the body opening into a collecting duct. The two collecting ducts typically meet up at a single bladder, opening to the exterior through one or two pores near the posterior end of the animal.[12]

The brain consists of a pair of ganglia in the head region, from which two or three pairs of nerve cords run down the length of the body. The nerve cords running along the ventral surface are always the largest, while the dorsal cords are present only in the Aspidogastrea. Trematodes generally lack any specialized sense organs, although some ectoparasitic species do possess one or two pairs of simple ocelli.[12]

Body wall musculature: Formed of three different muscle layers: circular, longitudinal, and diagonal. The outermost layer is formed by the circular muscle fibers, directly behind that are the longitudinal muscle fibers. The inner layer is formed by the diagonal muscle fibers. Together these muscle fibers form the segmented body wall of trematodes.[13]

Oral sucker and acetabulum: In some species of Trematoda, such as T. bragai, there is an acetabulum. This saucer-shaped organ is attached to the oral sucker in some Trematodes and other parasitic worms. This allows for parasitic worms to attach to their host by penetrating the host’s tissue with spines lining the acetabulum organ. In trematodes, the oral sucker is linked to the pharynx via a canal composed of meridional, equatorial, and radial muscle fibers.[13] Together, the mouth, pharynx, and esophagus form the foregut in Trematodes.[14]

Trematodes have a very complex life cycle and depending on what taxa they belong to, their life cycles can be completed with as little as one host compared to the typical three hosts.  When there is one host, this is normally a specific species of snail of the family Lymnaeidae. Almost all trematodes infect molluscs as the first host in the life cycle, and most have a complex life cycle involving other hosts. Most trematodes are monoecious and alternately reproduce sexually and asexually. The two main exceptions to this are the Aspidogastrea, which have no asexual reproduction, and the schistosomes, which are dioecious.

In the definitive host, in which sexual reproduction occurs, eggs are commonly shed along with host feces. Eggs shed in water release free-swimming larval forms (Miracidia) that are infective to the intermediate host, in which asexual reproduction occurs.

A species that exemplifies the remarkable life history of the trematodes is the bird fluke, Leucochloridium paradoxum. The definitive hosts, in which the parasite reproduces, are various woodland birds, while the hosts in which the parasite multiplies (intermediate host) are various species of snail. The adult parasite in the bird's gut produces eggs and these eventually end up on the ground in the bird's feces. Some eggs may be swallowed by a snail and hatch into larvae (miracidia). These larvae grow and take on a sac-like appearance. This stage is known as the sporocyst and it forms a central body in the snail's digestive gland that extends into a brood sac in the snail's head, muscular foot and eye-stalks. It is in the central body of the sporocyst where the parasite replicates itself, producing many tiny embryos (redia). These embryos move to the brood sac and mature into cercaria.

Trematodes can cause disease in many types of vertebrates, including mammals, birds, reptiles, and fish. Cattle and sheep can become infected by eating contaminated food. These infections lead to a reduction in milk or meat production, which can be of significant economic importance to the livestock industry.[14]

Human trematode infections are most common in Asia, Africa and Latin America. However, trematodes can be found anywhere where untreated human waste is used as fertilizer. Humans can be infected by trematodes by immersion in or ingestion of contaminated water, or by consuming raw or undercooked contaminated animals or plants.[16]

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