Schistosome
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''Schistosoma'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
trematode Trematoda is a Class (biology), class of flatworms known as trematodes, and commonly as flukes. They are obligate parasite, obligate Endoparasites, internal parasites with a complex biological life cycle, life cycle requiring at least two Host ( ...
s, commonly known as blood flukes. They are parasitic
flatworm Platyhelminthes (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") is a Phylum (biology), phylum of relatively simple bilaterian, Segmentation (biology), ...
s responsible for a highly significant group of
infection An infection is the invasion of tissue (biology), tissues by pathogens, their multiplication, and the reaction of host (biology), host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmis ...
s in
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s termed ''
schistosomiasis Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever is a neglected tropical helminthiasis, disease caused by parasitism, parasitic Schistosoma, flatworms called schistosomes. It affects both humans and animals. It affects ...
'', which is considered by the
World Health Organization The World Health Organization (WHO) is a list of specialized agencies of the United Nations, specialized agency of the United Nations which coordinates responses to international public health issues and emergencies. It is headquartered in Gen ...
to be the second-most socioeconomically devastating parasitic
disease A disease is a particular abnormal condition that adversely affects the structure or function (biology), function of all or part of an organism and is not immediately due to any external injury. Diseases are often known to be medical condi ...
(after
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
), infecting millions worldwide. Adult flatworms parasitize blood capillaries of either the mesenteries or plexus of the bladder, depending on the infecting species. They are unique among trematodes and any other flatworms in that they are
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
with distinct
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
between
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
and
female An organism's sex is female ( symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction. A female has larger gametes than a male. Females and ...
. Thousands of eggs are released and reach either the bladder or the intestine (according to the infecting species), and these are then excreted in
urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
or
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
to
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salt (chemistry), salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include ...
.
Larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e must then pass through an intermediate snail host before the next larval stage of the parasite emerges that can infect a new mammalian host by directly penetrating the skin.


Evolution

The origins of this
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
remain unclear. For many years it was believed that this genus had an African origin, but
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...
suggests that the
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
(''S. edwardiense'' and ''S. hippopotami'') that infect the hippo ('' Hippopotamus amphibius'') could be basal. Since hippos were present in both Africa and Asia during the
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
era, the genus might have originated as
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
of hippos. The original hosts for the South East Asian species were probably
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s. Based on the
phylogenetics In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
of the host snails it seems likely that the genus evolved in
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
between and . The
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to ''Schistosoma'' is a genus of
elephant Elephants are the largest living land animals. Three living species are currently recognised: the African bush elephant ('' Loxodonta africana''), the African forest elephant (''L. cyclotis''), and the Asian elephant ('' Elephas maximus ...
-infecting schistosomes — '' Bivitellobilharzia''. The
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
,
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
,
goat The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
and cashmere
goat The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
parasite '' Orientobilharzia turkestanicum'' appears to be related to the African schistosomes. This latter species has since been transferred to the genus ''Schistosoma''. Within the ''haematobium'' group ''S. bovis'' and ''S. curassoni'' appear to be closely related as do ''S. leiperi'' and ''S. mattheei''. ''S. mansoni'' appears to have evolved in
East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...
0.43–0.30 million years ago. ''S. mansoni'' and ''S. rodhaini'' appear to have shared a common ancestor between 107.5 and 147.6 thousand years ago. This period overlaps with the earliest archaeological evidence for fishing in Africa. It appears that ''S. mansoni'' originated in East Africa and experienced a decline in effective population size 20-90 thousand years ago before dispersing across the continent during the
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
. This species was later transmitted to the Americas by the slave trade. ''S. incognitum'' and ''S. nasale'' are more closely related to the African species rather than the ''japonicum'' group. ''S. sinensium'' appears to have radiated during the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
in the mid-Pleistocene. Estimated speciation dates for the ''japonicum'' group: ~3.8 million years ago for ''S. japonicum''/South East Asian schistosoma and ~2.5 million years ago for ''S. malayensis''/''S. mekongi''. ''Schistosoma turkestanicum'' is found infecting red deer in
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
. These strains appear to have diverged from those found in
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
and
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
. The date of divergence appears to be 270,000 years before present.


Taxonomy

The genus ''Schistosoma'' as currently defined is
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
, so revisions are likely. Over twenty species are recognised within this genus. The genus has been divided into four groups: ''indicum'', ''japonicum'', ''haematobium'' and ''mansoni''. The affinities of the remaining species are still being clarified. Thirteen species are found in Africa. Twelve of these are divided into two groups—those with a lateral spine on the egg (''mansoni'' group) and those with a terminal spine (''haematobium'' group).


Mansoni group

The four ''mansoni'' group species are: '' S. edwardiense'', '' S. hippotami'', '' S. mansoni'' and '' S. rodhaini''.


Haematobium group

The nine ''haematobium'' group species are: '' S. bovis'', '' S. curassoni'', '' S. guineensis'', '' S. haematobium'', '' S. intercalatum'', '' S. kisumuensis'', '' S. leiperi'', '' S. margrebowiei'' and '' S. mattheei''. ''S. leiperi'' and ''S. matthei'' appear to be related. ''S. margrebowiei'' is basal in this group. ''S. guineensis'' is the sister species to the ''S. bovis'' and ''S. curassoni'' grouping. ''S. intercalatum'' may actually be a species complex of at least two species.


Indicum group

The ''indicum'' group has three species: '' S. indicum'', '' S. nasale'' and '' S. spindale''. This group appears to have evolved during the Pleistocene. All use pulmonate
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s as hosts. ''S. spindale'' is widely distributed in Asia, Africa, and India.. ''S. indicum'' is found in India and
Thailand Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
. The indicum group appears to be the sister clade to the African species.


Japonicum group

The ''japonicum'' group has five species: '' S. japonicum'', '' S. malayensis'' and '' S. mekongi'', '' S. ovuncatum'' and '' S. sinensium'' and these species are found in
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
and Southeast Asia. '' S. ovuncatum'' forms a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
with '' S. sinensium'' and is found in northern Thailand. The definitive host is unknown and the intermediate host is the snail '' Tricula bollingi''. This species is known to use snails of the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Pomatiopsidae as hosts. '' S. incognitum'' appears to be basal in this genus. It may be more closely related to the African-Indian species than to the Southeast Asian group. This species uses pulmonate snails as hosts. Examination of the mitochondria suggests that ''Schistosoma incognitum'' may be a species complex.Webster BL, Littlewood DT (2012) Mitochondrial gene order change in ''Schistosoma'' (Platyhelminthes: Digenea: Schistosomatidae). Int J Parasitol 42(3):313-321


New species

As of 2012, four additional species have been transferred to this genus., previously classified as species in the genus ''Orientobilharzia''. Orientobilharzia differs from Schistosoma morphologically only on the basis of the number of testes. A review of the morphological and molecular data has shown that the differences between these genera are too small to justify their separation. The four species are *''Schistosoma bomfordi'' *''Schistosoma datta'' *''Schistosoma harinasutai'' *''Schistosoma turkestanicum''


Hybrids

The hybrid ''S. haematobium-S.guineenis'' was observed in Cameroon in 1996. ''S. haematobium'' could establish itself only after deforestation of the tropical rainforest in Loum next to the endemic ''S. guineensis''; hybridization led to competitive exclusion of ''S. guineensis''. In 2003, a ''S. mansoni-S. rodhaini'' hybrid was found in snails in western
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
, As of 2009, it had not been found in humans. In 2009, ''S. haematobium–S. bovis'' hybrids were described in northern Senegalese children. The Senegal River Basin had changed very much since the 1980s after the Diama Dam in Senegal and the Manantali Dam in Mali had been built. The Diama dam prevented ocean water to enter and allowed new forms of agriculture. Human migration, increasing number of livestock and sites where human and cattle both contaminate the water facilitated mixing between the different schistosomes in N'Der, for example. The same hybrid was identified during the 2015 investigation of a schistosomiasis outbreak on
Corsica Corsica ( , , ; ; ) is an island in the Mediterranean Sea and one of the Regions of France, 18 regions of France. It is the List of islands in the Mediterranean#By area, fourth-largest island in the Mediterranean and lies southeast of the Metro ...
, traced to the Cavu river. In 2019, a ''S. haematobium–S. mansoni'' hybrid was described in a 14-year-old patient with
hematuria Hematuria or haematuria is defined as the presence of blood or red blood cells in the urine. "Gross hematuria" occurs when urine appears red, brown, or tea-colored due to the presence of blood. Hematuria may also be subtle and only detectable with ...
from
Côte d'Ivoire Ivory Coast, also known as Côte d'Ivoire and officially the Republic of Côte d'Ivoire, is a country on the southern coast of West Africa. Its capital city of Yamoussoukro is located in the centre of the country, while its largest city and ...
.


Cladogram

A
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
based on
18S ribosomal RNA 18S ribosomal RNA (abbreviated 18S rRNA) is a part of the ribosomal RNA in eukaryotes. It is a component of the Eukaryotic small ribosomal subunit (40S) and the cytosolic homologue of both the 12S ribosomal RNA, 12S rRNA in mitochondria and the 1 ...
, 28S ribosomal RNA, and partial
cytochrome c oxidase subunit I Cytochrome c oxidase I (COX1) also known as mitochondrially encoded cytochrome c oxidase I (MT-CO1) is a protein that is encoded by the ''MT-CO1'' gene in eukaryotes. The gene is also called ''COX1'', ''CO1'', or ''COI''. Cytochrome c oxidase ...
(COI) genes shows phylogenic relations of species in the genus ''Schistosoma'':


Comparison of eggs

File:Schistosoma haematobium egg 4842 lores.jpg, '' Schistosoma japonicum'' File:Schistosoma japonicum egg 4843 lores.jpg, '' Schistosoma haematobium'' File:Schistosoma mansoni egg 4841 lores.jpg, '' Schistosoma mansoni'' File:S mekongi eggR.jpg, '' Schistosoma mekongi'' File:S interculatum eggS.jpg, '' Schistosoma intercalatum''


Geographical distribution

Geographical areas associated with schistosomiasis by the World Health Organization as of January 2017 include in alphabetical order: Africa, Brazil, Cambodia, the Caribbean, China, Corsica, Indonesia, Laos, the Middle East, the Philippines, Suriname, and Venezuela. There had been no cases in Europe since 1965, until an outbreak occurred on Corsica.


Schistosomiasis

The parasitic flatworms of ''Schistosoma'' cause a group of chronic infections called
schistosomiasis Schistosomiasis, also known as snail fever, bilharzia, and Katayama fever is a neglected tropical helminthiasis, disease caused by parasitism, parasitic Schistosoma, flatworms called schistosomes. It affects both humans and animals. It affects ...
known also as bilharziasis. An anti-schistosome drug is a schistosomicide.


Species infecting humans

Parasitism of humans by ''Schistosoma'' appears to have evolved at least three occasions in both
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
and
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
. *'' S. guineensis'', a recently described species, is found in
West Africa West Africa, also known as Western Africa, is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Gha ...
. Known
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
intermediate hosts include '' Bulinus forskalii.'' *'' S. haematobium'', commonly referred to as the ''bladder fluke'', originally found in Africa, the
Near East The Near East () is a transcontinental region around the Eastern Mediterranean encompassing the historical Fertile Crescent, the Levant, Anatolia, Egypt, Mesopotamia, and coastal areas of the Arabian Peninsula. The term was invented in the 20th ...
, and the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
basin, was introduced into India during World War II. Freshwater snails of the genus '' Bulinus'' are an important intermediate host for this parasite. Among final hosts humans are most important. Other final hosts are rarely baboons and monkeys. *'' S. intercalatum.'' The usual final hosts are humans. Other animals can be infected experimentally. *'' S. japonicum'', whose common name is simply ''blood fluke'', is widespread in
East Asia East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
and the southwestern
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is bounded by the cont ...
region. Freshwater snails of the genus '' Oncomelania'' are an important intermediate host for ''S. japonicum.'' Final hosts are humans and other mammals including cats, dogs, goats, horses, pigs, rats and water buffalo. *'' S. malayensis'' This species appears to be a rare infection in humans and is considered to be a
zoonosis A zoonosis (; plural zoonoses) or zoonotic disease is an infectious disease of humans caused by a pathogen (an infectious agent, such as a virus, bacterium, parasite, fungi, or prion) that can jump from a non-human vertebrate to a human. When ...
. The natural vertebrate host is Müller's giant Sunda rat (''Sundamys muelleri''). The snail hosts are Robertsiella species (''R. gismanni'', ''R. kaporensis'' and ''R. silvicola'' (see Attwood et al. 2005 Journal of Molluscan Studies Volume 71, Issue 4 pp. 379–391). *'' S. mansoni'', found in
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
,
Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...
,
Suriname Suriname, officially the Republic of Suriname, is a country in northern South America, also considered as part of the Caribbean and the West Indies. It is a developing country with a Human Development Index, high level of human development; i ...
, the lesser
Antilles The Antilles is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mexico to the northwest, and the Atlantic Ocean to the north and east. The Antillean islands are divided into two smaller groupings: the Greater An ...
,
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
, and the
Dominican Republic The Dominican Republic is a country located on the island of Hispaniola in the Greater Antilles of the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean. It shares a Maritime boundary, maritime border with Puerto Rico to the east and ...
. It is also known as ''Manson's blood fluke'' or ''swamp fever''. Freshwater snails of the genus '' Biomphalaria'' are an important intermediate host for this trematode. Among final hosts humans are most important. Other final hosts are baboons, rodents and raccoons. *'' S. mekongi'' is related to ''S. japonicum'' and affects both the superior and inferior mesenteric veins. ''S. mekongi'' differs in that it has smaller eggs, a different intermediate host ('' Neotricula aperta'') and longer prepatent period in the mammalian host. Final hosts are humans and dogs. The snail '' Tricula aperta'' can also be experimentally infected with this species.


Species infecting other animals

'' Schistosoma indicum'', '' Schistosoma nasale'', '' Schistosoma spindale'', '' Schistosoma leiperi'' are all parasites of
ruminant Ruminants are herbivorous grazing or browsing artiodactyls belonging to the suborder Ruminantia that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microb ...
s. '' Schistosoma edwardiense'' and '' Schistosoma hippopotami'' are parasites of the hippo. '' Schistosoma ovuncatum'' and '' Schistosoma sinensium'' are parasites of rodents.


Morphology

Adult schistosomes share all the fundamental features of the digenea. They have a basic
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symme ...
, oral and ventral suckers, a body covering of a syncytial tegument, a blind-ending
digestive system The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
consisting of
mouth A mouth also referred to as the oral is the body orifice through which many animals ingest food and animal communication#Auditory, vocalize. The body cavity immediately behind the mouth opening, known as the oral cavity (or in Latin), is also t ...
,
esophagus The esophagus (American English), oesophagus (British English), or œsophagus (Œ, archaic spelling) (American and British English spelling differences#ae and oe, see spelling difference) all ; : ((o)e)(œ)sophagi or ((o)e)(œ)sophaguses), c ...
and bifurcated caeca; the area between the tegument and alimentary canal filled with a loose network of
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical ...
cells, and an excretory or osmoregulatory system based on
flame cell A flame cell is a specialized excretory cell found in simple invertebrates, including flatworms ( Platyhelminthes), rotifers and nemerteans; these are the simplest animals to have a dedicated excretory system. Flame cells function like a kidney ...
s. Adult worms tend to be long and use
globin The globins are a superfamily of heme-containing globular proteins, involved in binding and/or transporting oxygen. These proteins all incorporate the globin fold, a series of eight alpha helical segments. Two prominent members include myo ...
s from their hosts'
hemoglobin Hemoglobin (haemoglobin, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobin ...
for their own circulatory system.


Reproduction

Unlike other trematodes and basically all other flatworms, the schistosomes are
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, ''i.e.'', the sexes are separate. The two sexes display a strong degree of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
, and the male is considerably larger than the female. The male surrounds the female and encloses her within his ''gynacophoric canal'' for the entire adult lives of the worms. As the male feeds on the host's blood, he passes some of it to the female. The male also passes on chemicals which complete the female's development, whereupon they will reproduce sexually. Although rare, sometimes mated schistosomes will "divorce", wherein the female will leave the male for another male. The exact reason is not understood, although it is thought that females will leave their partners to mate with more genetically distant males. Such a biological mechanism would serve to decrease inbreeding, and may be a factor behind the unusually high genetic diversity of schistosomes.


Genome

The genomes of ''Schistosoma haematobium'', ''S. japonicum'' and '' S. mansoni ''have been reported.


History

The eggs of these
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
were first seen by Theodor Maximilian Bilharz, a German
pathologist Pathology is the study of disease. The word ''pathology'' also refers to the study of disease in general, incorporating a wide range of biology research fields and medical practices. However, when used in the context of modern medical treatme ...
working in
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
in 1851 who found the eggs of '' Schistosoma haematobium'' during the course of a post mortem. He wrote two letters to his former teacher von Siebold in May and August 1851 describing his findings. Von Siebold published a paper in 1852 summarizing Bilharz's findings and naming the worms ''Distoma haematobium''. Bilharz wrote a paper in 1856 describing the
worm Worms are many different distantly related bilateria, bilateral animals that typically have a long cylindrical tube-like body, no limb (anatomy), limbs, and usually no eyes. Worms vary in size from microscopic to over in length for marine ...
s more fully. Their unusual morphology meant that they could not be comfortably included in ''Distoma''. So in 1856 Meckel von Helmsback ( de) created the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''Bilharzia'' for them. In 1858 David Friedrich Weinland proposed the name ''Schistosoma'' (Greek: "split body") because the worms were not hermaphroditic but had separate sexes. Despite ''Bilharzia'' having precedence, the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
name ''Schistosoma'' was officially adopted by the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
. The term ''Bilharzia'' to describe infection with these parasites is still in use in medical circles. Bilharz also described '' Schistosoma mansoni'', but this species was redescribed by Louis Westenra Sambon in 1907 at the London School of Tropical Medicine who named it after his teacher Patrick Manson. In 1898, all then known species were placed in a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
by Stiles and Hassel. This was elevated to family status by Looss in 1899. Poche in 1907 corrected a grammatical error in the family name. The life cycle of ''Schistosoma mansoni'' was determined by the Brazilian parasitologist Pirajá da Silva (1873-1961) in 1908. In 2009, the genomes of ''Schistosoma mansoni'' and '' Schistosoma japonicum'' were decoded opening the way for new targeted treatments. In particular, the study discovered that the genome of ''S. mansoni'' contained 11,809
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
s, including many that produce
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s for breaking down
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s, enabling the parasite to bore through tissue. Also, ''S. mansoni'' does not have an enzyme to make certain
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...
s, so it must rely on its host to produce these.


Treatment

Praziquantel


References


Further reading

*


External links


British Department for International Development Control of SchistosomiasisThe World Health Organisation page on SchistosomiasisUniversity of Cambridge Schistosome LaboratorySchistosoma parasites overview, biology, life cycle image at MetaPathogen
{{Authority control Digenea genera Parasitic helminths of humans Diplostomida