Product Code Database
Platyhelminthes (from the Greek language πλατύ, platy, meaning "flat" and ἕλμινς (root: ἑλμινθ-), helminth-, meaning "worm") is a phylum of relatively simple bilaterian, unsegmented, soft-bodied commonly called flatworms or flat worms. Being (having no coelom), and having no specialised circulatory and respiratory organs, they are restricted to having flattened shapes that allow oxygen and nutrients to pass through their bodies by diffusion. The digestive cavity has only one opening for both ingestion (intake of nutrients) and egestion (removal of undigested wastes); as a result, the food can not be processed continuously.
In traditional medicinal texts, Platyhelminthes are divided into Turbellaria, which are mostly non-parasitic such as , and three entirely parasitic groups: Cestoda, Trematoda and Monogenea; however, since the turbellarians have since been proven not to be monophyletic, this classification is now deprecated. Free-living flatworms are mostly predators, and live in water or in shaded, humid terrestrial environments, such as leaf litter. Cestodes (tapeworms) and trematodes (flukes) have complex life-cycles, with mature stages that live as parasites in the digestive systems of fish or land , and intermediate stages that infest secondary hosts. The eggs of trematodes are excreted from their main hosts, whereas adult cestodes generate vast numbers of hermaphroditic, segment-like that detach when mature, are excreted, and then release eggs. Unlike the other parasitic groups, the monogeneans are external parasites infesting , and their metamorphose into the adult form after attaching to a suitable host.
Because they do not have internal body Body cavity, Platyhelminthes were regarded as a primitive stage in the evolution of bilaterians (animals with bilateral symmetry and hence with distinct front and rear ends). However, analyses since the mid-1980s have separated out one subgroup, the Acoelomorpha, as basal bilaterians – closer to the original than to any other modern groups. The remaining Platyhelminthes form a monophyletic group, one that contains all and only descendants of a common ancestor that is itself a member of the group. The redefined Platyhelminthes is part of the Spiralia, one of the two main groups of Protostomia. These analyses had concluded the redefined Platyhelminthes, excluding Acoelomorpha, consists of two monophyletic subgroups, Catenulida and Rhabditophora, with Cestoda, Trematoda and Monogenea forming a monophyletic subgroup within one branch of the Rhabditophora. Hence, the traditional platyhelminth subgroup "Turbellaria" is now regarded as paraphyletic, since it excludes the wholly parasitic groups, although these are descended from one group of "turbellarians".
A planarian species has been used in the Philippines and Maldives in an attempt to control populations of the imported giant African snail ( Achatina fulica), which was eating agricultural crops. Success was initially reported for the Maldives but this was only temporary and the role of flatworms has been questioned. These planarians have now spread very widely throughout the tropics and are themselves a serious threat to native snails, and should not be used for biological control. In northwest Europe, there are concerns about the spread of the New Zealand planarian Arthurdendyus triangulatus, which preys on .
The space between the skin and gut is filled with mesenchyme, also known as parenchyma, a connective tissue made of cells and reinforced by collagen fibers that act as a type of skeleton, providing attachment points for . The mesenchyme contains all the internal organs and allows the passage of oxygen, nutrients and waste products. It consists of two main types of cell: fixed cells, some of which have fluid-filled ; and , which can transform into any other type of cell, and are used in regenerating tissues after injury or asexual reproduction.
Most platyhelminths have no anus and regurgitate undigested material through the mouth. The genus Paracatenula, whose members include tiny flatworms living in symbiosis with bacteria, is even missing a mouth and a gut. However, some long species have an anus and some with complex, branched guts have more than one anus, since excretion only through the mouth would be difficult for them. The gut is lined with a single layer of cells that absorb and digest food. Some species break up and soften food first by secreting in the gut or pharynx (throat).
All animals need to keep the concentration of dissolved substances in their body fluids at a fairly constant level. Internal parasites and free-living marine animals live in environments with high concentrations of dissolved material, and generally let their tissues have the same level of concentration as the environment, while freshwater animals need to prevent their body fluids from becoming too dilute. Despite this difference in environments, most platyhelminths use the same system to osmoregulation of their body fluids. , so called because the beating of their flagella looks like a flickering candle flame, extract from the mesenchyme water that contains wastes and some reusable material, and drive it into networks of tube cells which are lined with flagella and microvilli. The tube cells' flagella drive the water towards exits called , while their microvilli reabsorb reusable materials and as much water as is needed to keep the body fluids at the right concentration. These combinations of flame cells and tube cells are called protonephridia.
In all platyhelminths, the nervous system is concentrated at the head end. Other platyhelminths have rings of Ganglion in the head and main nerve trunks running along their bodies.
Some turbellarians have a simple pharynx lined with Cilium and generally feed by using cilia to sweep food particles and small prey into their mouths, which are usually in the middle of their undersides. Most other turbellarians have a pharynx that is eversible (can be extended by being turned inside-out), and the mouths of different species can be anywhere along the underside. The freshwater species Microstomum caudatum can open its mouth almost as wide as its body is long, to swallow prey about as large as itself. Predatory species in suborder Kalyptorhynchia often have a muscular pharynx equipped with hooks or teeth used for seizing prey.
Most turbellarians have pigment-cup ocelli ("little eyes"); one pair in most species, but two or even three pairs in others. A few large species have many eyes in clusters over the brain, mounted on tentacles, or spaced uniformly around the edge of the body. The ocelli can only distinguish the direction from which light is coming to enable the animals to avoid it. A few groups have - fluid-filled chambers containing a small, solid particle or, in a few groups, two. These statocysts are thought to function as balance and acceleration sensors, as they perform the same way in medusae and in . However, turbellarian statocysts have no sensory cilia, so the way they sense the movements and positions of solid particles is unknown. On the other hand, most have ciliated touch-sensor cells scattered over their bodies, especially on tentacles and around the edges. Specialized cells in pits or grooves on the head are most likely smell sensors.
, a subgroup of seriates, are famous for their ability to regenerate if divided by cuts across their bodies. Experiments show that (in fragments that do not already have a head) a new head grows most quickly on those fragments which were originally located closest to the original head. This suggests the growth of a head is controlled by a chemical whose concentration diminishes throughout the organism, from head to tail. Many turbellarians cloning themselves by transverse or longitudinal division, whilst others, reproduce by budding.
The vast majority of turbellarians are (they have both female and male reproductive cells) which fertilize eggs internally by copulation. Some of the larger aquatic species mate by penis fencing – a duel in which each tries to impregnate the other, and the loser adopts the female role of developing the eggs. In most species, "miniature adults" emerge when the eggs hatch, but a few large species produce plankton-like .
Adults range between and in length. Individual adult digeneans are of a single sex, and in some species slender females live in enclosed grooves that run along the bodies of the males, partially emerging to lay eggs. In all species the adults have complex reproductive systems, capable of producing between 10,000 and 100,000 times as many eggs as a free-living flatworm. In addition, the intermediate stages that live in snails reproduce asexually.
Adults of different species infest different parts of the definitive host - for example the intestine, , large blood vessels, and liver. The adults use a relatively large, muscular pharynx to ingest cells, cell fragments, mucus, body fluids or blood. In both the adult and snail-inhabiting stages, the external syncytium absorbs dissolved nutrients from the host. Adult digeneans can live without oxygen for long periods.
In the majority of species, known as eucestodes ("true tapeworms"), the neck produces a chain of segments called proglottids via a process known as strobilation. As a result, the most mature proglottids are furthest from the scolex. Adults of Taenia saginata, which infests humans, can form proglottid chains over long, although is more typical. Each proglottid has both male and female reproductive organs. If the host's gut contains two or more adults of the same cestode species they generally fertilize each other, however, proglottids of the same worm can fertilize each other and even themselves. When the eggs are fully developed, the proglottids separate and are excreted by the host. The eucestode life cycle is less complex than that of , but varies depending on the species. For example:
Members of the smaller group known as Cestodaria have no scolex, do not produce proglottids, and have body shapes similar to those of diageneans. Cestodarians parasitize fish and turtles.
The internal relationships of Platyhelminthes are shown below. The tree is not fully resolved.
The oldest confidently identified parasitic flatworm fossils are cestode eggs found in a Permian shark coprolite, but helminth hooks still attached to Devonian acanthodians and placoderms might also represent parasitic flatworms with simple life cycles.De Baets, K., P. Dentzien-Dias, I. Upeniece, O. Verneau and P. C. J. Donoghue (2015). 9780128040010 . ISBN 9780128040010 The oldest known free-living platyhelminth specimen is a fossil preserved in Eocene age Baltic amber and placed in the monotypic species Micropalaeosoma, whilst the oldest subfossil specimens are Schistosoma eggs discovered in ancient Egyptian Mummy. The Platyhelminthes have very few synapomorphy - distinguishing features that all Platyhelminthes (but no other animals) exhibit. This makes it difficult to work out their relationships with other groups of animals, as well as the relationships between different groups that are described as members of the Platyhelminthes.
The "traditional" view before the 1990s was that Platyhelminthes formed the sister group to all the other bilaterians, which include, for instance, , , and . Since then, molecular phylogenetics, which aims to work out evolutionary "family trees" by comparing different organisms' biochemistry such as DNA, RNA and , has radically changed scientists' view of evolutionary relationships between animals. Flatworms are now recognized as secondarily simplified bilaterians.
Detailed morphological analyses of anatomical features in the mid-1980s, as well as molecular phylogenetics analyses since 2000 using different sections of DNA, agree that Acoelomorpha, consisting of Acoela (traditionally regarded as very simple "") and Nemertodermatida (another small group previously classified as "turbellarians") are the sister group to all other bilaterians. However, a 2007 study concluded that Acoela and Nemertodermatida were two distinct groups of bilaterians.
Xenoturbella, a bilaterian whose only well-defined organ is a statocyst, was originally classified as a "primitive turbellarian". Later studies suggested it may instead be a deuterostome, but more detailed molecular phylogenetics have led to its classification as sister-group to the Acoelomorpha.
The Platyhelminthes excluding Acoelomorpha contain two main groups - Catenulida and Rhabditophora - both of which are generally agreed to be monophyletic (each contains all and only the descendants of an ancestor that is a member of the same group). (2025). 9780195172348, Oxford University Press US. ISBN 9780195172348 Early molecular phylogenetics analyses of the Catenulida and Rhabditophora left uncertainties about whether these could be combined in a single monophyletic group; a study in 2008 concluded that they could, therefore Platyhelminthes could be redefined as Catenulida plus Rhabditophora, excluding the Acoelomorpha.
Other molecular phylogenetics analyses agree the redefined Platyhelminthes are most closely related to Gastrotricha, and both are part of a grouping known as Platyzoa. Platyzoa are generally agreed to be at least closely related to the Lophotrochozoa, a superphylum that includes molluscs and annelid worms. The majority view is that Platyzoa are part of Lophotrochozoa, but a significant minority of researchers regard Platyzoa as a sister group of Lophotrochozoa.
It has been agreed since 1985 that each of the wholly parasitic platyhelminth groups (Cestoda, Monogenea and Trematoda) is monophyletic, and that together these form a larger monophyletic grouping, the Neodermata, in which the adults of all members have syncytium skins. However, there is debate about whether the Cestoda and Monogenea can be combined as an intermediate monophyletic group, the Cercomeromorpha, within the Neodermata. It is generally agreed that the Neodermata are a sub-group a few levels down in the "family tree" of the Rhabditophora. Hence the traditional sub-phylum "Turbellaria" is paraphyletic, since it does not include the Neodermata although these are descendants of a sub-group of "turbellarians".
In several members of the order Rhabdocoela an Endosymbiont relationship with microalgae has evolved. Some species in the same order has also evolved kleptoplasty.
The earliest known fossils confidently classified as tapeworms have been dated to , after being found in (fossilised faeces) from an elasmobranch. Putative older fossils include a ribbon-shaped, bilaterally symmetrical organism named Rugosusivitta orthogonia from the Early Cambrian of China, brownish bodies on the bedding planes reported from the Late Ordovician (Katian) Vauréal Formation (Canada) by Knaust & Desrochers (2019), tentatively interpreted as turbellarians (though the authors cautioned that they might ultimately turn out to be fossils of Acoelomorpha or ) and circlets of fossil hooks preserved with Placodermi and Acanthodii fossils from the Devonian of Latvia, at least some of which might represent parasitic monogeneans.
In 2000, an estimated 45 million people were infected with the beef tapeworm Taenia saginata and 3 million with the pork tapeworm Taenia solium. Infection of the digestive system by adult tapeworms causes abdominal symptoms that, whilst unpleasant, are seldom disabling or life-threatening. However, neurocysticercosis resulting from penetration of T. solium larvae into the central nervous system is the major cause of acquired epilepsy worldwide. In 2000, about 39 million people were infected with (flukes) that naturally parasitize fish and crustaceans, but can pass to humans who eat raw or lightly cooked seafood. Infection of humans by the broad fish tapeworm Diphyllobothrium latum occasionally causes vitamin B12 deficiency and, in severe cases, megaloblastic anemia.
The threat to humans in developed countries is rising as a result of social trends: the increase in organic farming, which uses manure and sewage sludge rather than artificial , spreads parasites both directly and via the droppings of which feed on manure and sludge; the increasing popularity of raw or lightly cooked foods; imports of meat, seafood and salad vegetables from high-risk areas; and, as an underlying cause, reduced awareness of parasites compared with other public health issues such as pollution. In less-developed countries, inadequate sanitation and the use of human feces (night soil) as fertilizer or to enrich fish farm ponds continues to spread parasitic platyhelminths, whilst poorly designed water-supply and irrigation projects have provided additional channels for their spread. People in these countries usually cannot afford the cost of fuel required to cook food thoroughly enough to kill parasites. Controlling parasites that infect humans and livestock has become more difficult, as many species have become Drug resistance to drugs that used to be effective, mainly for killing juveniles in meat. While poorer countries still struggle with unintentional infection, cases have been reported of intentional infection in the US by dieters who are desperate for rapid weight-loss.
A study in Argentina shows the potential for planarians such as Girardia anceps, Mesostoma ehrenbergii, and Bothromesostoma evelinae to reduce populations of the mosquito species Aedes aegypti and Culex pipiens. The experiment showed that G. anceps can prey on all instars of both mosquito species, yet maintain a steady predation rate over time. The ability of these flatworms to live in artificial containers demonstrated the potential of placing these species in popular mosquito breeding sites, which might reduce the amount of mosquito-borne disease.
|
|
