''Sarcocystis'' 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
protozoa
Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
n
parasite
Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
s, with many species infecting
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s,
reptile
Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s and
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s. Its name is derived from
Greek
Greek may refer to:
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group
*Greek language, a branch of the Indo-European language family
**Proto-Greek language, the assumed last common ancestor of all kno ...
''sarx'' = flesh and ''kystis'' = bladder.
The lifecycle of a typical member of this genus involves two host species, a definitive host and an intermediate host. Often, the definitive host is a predator and the intermediate host is its prey. The parasite reproduces sexually in the gut of the definitive host, is passed with the feces, and ingested by the intermediate host. There, it eventually enters muscle tissue. When the intermediate host is eaten by the definitive host, the cycle is completed. The definitive host usually does not show any symptoms of infection, but the intermediate host does.
About 130 recognized
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), ...
are in this genus. Revision of the taxonomy of the genus is ongoing, and all the currently recognized species may be a much smaller number of species that can infect multiple hosts.
History
The organism was first recognized in a
mouse
A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus'' ...
by Miescher in 1843.
His findings were not initially interpreted as involving a protist, and the literature referred to the structures he described as "Miescher's tubules". Incidentally, Miescher's son,
Johann Friedrich Miescher, discovered
DNA
Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
. Similar structures were found in
pig
The pig (''Sus domesticus''), also called swine (: swine) or hog, is an omnivorous, domesticated, even-toed, hoofed mammal. It is named the domestic pig when distinguishing it from other members of the genus '' Sus''. Some authorities cons ...
muscle in 1865, but these remained unnamed until 1899, when the name ''Sarcocystis miescheriana'' was proposed for them.
Initially, whether these organisms were
fungi
A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
or protozoa was unclear . This uncertainty was resolved in 1967 when electron microscopic studies showed that they were protozoa, related to ''
Toxoplasma'' and ''
Eimeria
''Eimeria'' is a genus of apicomplexan parasites that includes various species capable of causing the disease coccidiosis in animals such as cattle, poultry and smaller ruminants including sheep and goats. ''Eimeria'' species are considered to ...
''. The lifecycle remained unknown until 1970, when bradyzoites from sarcocysts in bird muscles were inoculated into cultured mammalian cells and seen to undergo development into sexual stages and
oocysts. Transmission studies with ''Sarcocystis'' of cattle (then considered a single species, ''Sarcocystis fusiformis'') in dogs, cats, and humans revealed three morphologically distinct species, which were named ''S. bovicanis'', ''S. bovifelis'', and ''S. bovihominis''. This and post-1972 research on ''Sarcocystis'' was reviewed during the same decade; and that account is still a very useful source of information today.
Lifecycle
The
heteroxenous
Heteroxeny, or heteroxenous development, characterizes a parasite whose development involves several host species. Heteroxeny has been used as the basis for splitting genera.
When there are two or three hosts, the development cycle is named d ...
(more than one obligatory host) lifecycle of these
apicomplexan parasite
Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
s remained obscure until 1972, when the prey-predator relationship of its definitive and intermediate hosts was recognized. The lifecycles of about 60 of these species are now known.
In outline,
gametogony and
sporogony occur in the
intestine
The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascular system. T ...
of the definitive host, while both
schizogony, which occurs in various tissues, and the formation of
sarcocysts (containing
bradyzoites and
metrocytes) occurs principally in the
muscle
Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
s of the intermediate host. In some cases, a single species may act as both the definitive and intermediate host.
Oocysts
Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism ...
are passed in the
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 ...
of an infected definitive host. The oocyst undergoes sporogony, creating two
sporocysts. These sporocysts of ''Sarcocystis'' characteristically contain four
sporozoite
Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organis ...
s and measure approximately 15–19 by 8–10 μm. Oocysts of ''Sarcocystis'' are thin-walled and easily break open. The result is that sporocysts recovered from feces outnumber intact oocysts.
Intermediate hosts such as
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 ...
or pigs then ingest sporocysts. Sporozoites are released in the body and migrate to vessels, where they undergo the first two generations of asexual reproduction. These rounds result in the development of
meronts. This stage lasts about 15 to 16 days after ingestion of sporocysts.
Merozoites
Apicomplexans, a group of intracellular parasites, have life cycle stages that allow them to survive the wide variety of environments they are exposed to during their complex life cycle. Each stage in the life cycle of an apicomplexan organism ...
emerge from the second-generation meronts and enter the mononucleated cells, where they develop by
endodyogeny. Subsequent generations of merozoites develop downstream in the direction of blood flow to arterioles, capillaries, venules, and veins throughout the body, subsequently developing into the final asexual generation in muscles.
Merozoites entering muscle cells round up to form
metrocytes and initiate
sarcocyst formation. Sarcocysts begin as unicellular bodies containing a single metrocyte and through asexual multiplication numerous metrocytes accumulate and the sarcocyst increases in size. As the sarcocyst matures, the small, rounded, noninfectious metrocytes give rise to crescent-shaped bodies called
bradyzoites (also known as "bradyzoic merozoites") that are infectious for the definitive host. The time required for maturation varies with the species and may take 2 months or more.
In species in which symptoms develop, these typically occur 20–40 days after ingestion of sporocysts and during the subsequent migration of sporozoites through the body vessels. Acute lesions (oedema, hemorrhages, and necrosis) develop in the affected tissues. The parasite has a predilection for skeletal muscle (myositis), cardiac muscle (petechial hemorrhages of cardiac muscle and serosae), and lymph nodes (oedema, necrosis, and hemorrhage). These lesions are associated with maturation of second generation of meronts within the endothelial and subendothelials cells. Occasionally mononuclear infiltration or hyperemia has been observed in the lamina propria of the small intestine. After the acute phase, cysts may be found in various muscular tissues, generally without pathology.
Once the intermediate host is eaten by the definitive host, such as a dog or human, the parasite undergoes sexual reproduction within the gut to create
macrogamonts and
microgamonts. Most definitive hosts do not show any clinical signs or symptoms. Fusion of a macrogamont and a microgamont creates a
zygote
A zygote (; , ) is a eukaryote, eukaryotic cell (biology), cell formed by a fertilization event between two gametes.
The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individ ...
, which develops into an oocyst. The oocyst is passed through the faeces, completing the lifecycle.
A second lifecycle has more recently been described whereby
carnivore
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s and
omnivore
An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...
s pass the infectious stages in their faeces. Ingestion of this material may lead to successful infection of the ingesting animal.
Birds
Although sarcocysts were first reported in the muscles of birds by Kuhn in 1865, the first lifecycle involving a bird (''
Gallus gallus
The red junglefowl (''Gallus gallus''), also known as the Indian red junglefowl (and formerly the bankiva or bankiva-fowl), is a species of Tropics, tropical, predominantly Terrestrial animal, terrestrial bird in the fowl and pheasant family, P ...
'') and a carnivore (''
Canis familiaris
The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a Domestication of vertebrates, domesticated descendant of the gray wolf. Also called the domestic dog, it was Selective breeding, selectively bred from a population of wolves ...
'') was not described until 1977 by Munday et al.
In 1986 the first life cycle involving birds as both the definitive (northern goshawk – ''
Accipiter gentilis'') and intermediate (Atlantic canary – ''
Serinus canaria'') hosts was described by Cerná and Kvasnovská.
Taxonomy
The taxonomy of this genus and its relationship to other protozoal genera are currently under investigation.
Related genera include: ''
Besnoitia'', ''
Caryospora'', ''
Cystoisospora'', ''
Frenkelia'', ''
Isospora'', ''
Hammondia'', ''
Hyaloklossia'', ''
Lankesterella'', ''
Neospora'', and ''
Toxoplasma''.
''Sarcocystis'' is the largest genus within the family Sarcocystidae and consists of species that infect a range of animals, including mammals, birds, and reptiles. ''Frenkelia'', another genus within this family, consists of parasites that use rodents as intermediate hosts and birds of prey as definitive hosts.
''Besnoitia'', ''Hammondia'', ''Neospora'', and ''Toxoplasma'' apparently form a single
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 ...
. Within this clade, ''Toxoplasma'' and ''Neospora'' appear to be sister clades. ''Isospora'' also appears to belong to this clade and this clade is a sister to ''Sarcocystis''. ''Frenkelia'' appears to be very closely related to ''Sarcocystis''.
Several molecular studies have suggested that ''Frenkelia'' is actually a species of ''Sarcocystis''. This genus was distinguished from ''Sarcocystis'' on the basis of its tendency to encyst within the
brain
The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
rather than within muscle. This distinction may not be taxonomically valid.
Within the genus, a number of clades have been identified. These include one that contains ''S. dispersa'', ''S. lacertae'', ''S. mucosa'', ''S. muris'', ''S. neurona'', and ''S. rodentifelis''.
''Frenkelia'' also groups with this clade.
Species
* ''
Sarcocystis accipitris''
* ''
Sarcocystis alces''
* ''
Sarcocystis alceslatrans''
* ''
Sarcocystis ameivamastigodryasi''
* ''
Sarcocystis americana''
* ''
Sarcocystis arieticanis''
* ''
Sarcocystis asinus''
* ''
Sarcocystis atheridis''
* ''
Sarcocystis aucheniae''
* ''
Sarcocystis bertrami''
* ''
Sarcocystis bigemina''
* ''
Sarcocystis booliati''
* ''
Sarcocystis bovicanis''
* ''
Sarcocystis bovifelis''
* ''
Sarcocystis bovihominis''
* ''
Sarcocystis buffalonis''
* ''
Sarcocystis calchasi''
* ''
Sarcocystis cameli''
* ''
Sarcocystis camelopardalis''
* ''
Sarcocystis campestris''
* ''
Sarcocystis chamaleonis''
* ''
Sarcocystis cernae''
* ''
Sarcocystis cervi''
* ''
Sarcocystis cervicanis''
* ''
Sarcocystis canis''
* ''
Sarcocystis capracanis''
* ''
Sarcocystis cornixi''
* ''
Sarcocystis crotali''
* ''
Sarcocystis cruzi''
* ''
Sarcocystis cuniculi''
* ''
Sarcocystis cymruensis''
* ''
Sarcocystis danzani''
* ''
Sarcocystis dasypi''
* ''
Sarcocystis debonei''
* ''
Sarcocystis diminuta''
* ''
Sarcocystis dirumpens''
* ''
Sarcocystis dispersa''
* ''
Sarcocystis dubeyella''
* ''
Sarcocystis dubeyi''
* ''
Sarcocystis equicanis''
* ''
Sarcocystis falcatula''
* ''
Sarcocystis fayeri''
* ''
Sarcocystis felis''
* ''
Sarcocystis fulicae''
* ''
Sarcocystis fusiformis''
* ''
Sarcocystis gallotiae''
* ''
Sarcocystis garnhami''
* ''
Sarcocystis gracilis''
* ''
Sarcocystis grueneri''
* ''
Sarcocystis halieti''
* ''
Sarcocystis hoarensis''
* ''
Sarcocystis fusiformis''
* ''
Sarcocystis gallotiae''
* ''
Sarcocystis gerbilliechis''
* ''
Sarcocystis gigantea''
* ''
Sarcocystis giraffae''
* ''
Sarcocystis gongyli''
* ''
Sarcocystis gracilis''
* ''
Sarcocystis greineri''
* ''
Sarcocystis grueneri''
* ''
Sarcocystis hardangeri''
* ''
Sarcocystis hemioni''
* ''
Sarcocystis hemionilatrantis''
* ''
Sarcocystis hericanis''
* ''
Sarcocystis hircicanis''
* ''
Sarcocystis hirsuta''
* ''
Sarcocystis hjorti''
* ''
Sarcocystis hofmanni''
* ''
Sarcocystis hominis''
* ''
Sarcocystis horvathi''
* ''
Sarcocystis iberica''
* ''
Sarcocystis idahoensis''
* ''
Sarcocystis inghami''
* ''
Sarcocystis jamaicensis''
* ''
Sarcocystis jorrini''
* ''
Sarcocystis kinosterni''
* ''
Sarcocystis kirkpatricki''
* ''
Sarcocystis kirmsei''
[ ]
* ''
Sarcocystis klaseriensis''
* ''
Sarcocystis kortei''
* ''
Sarcocystis lacertae''
* ''
Sarcocystis lamacanis''
* ''
Sarcocystis leporum''
* ''
Sarcocystis levinei''
* ''
Sarcocystis lindemanni''
* ''
Sarcocystis lindsayi''
* ''
Sarcocystis linearis''
* ''Sarcocystis markusi''
* ''
Sarcocystis medusiformis''
* ''
Sarcocystis melis''
* ''
Sarcocystis mephitisi''
* ''
Sarcocystis miescheriana''
* ''
Sarcocystis mihoensis''
* ''
Sarcocystis mitrani''
* ''
Sarcocystis mongolica''
* ''
Sarcocystis montanaensis''
* ''
Sarcocystis morae''
* ''
Sarcocystis mucosa''
* ''
Sarcocystis moulei''
* ''
Sarcocystis murinotechis''
* ''
Sarcocystis muris''
* ''
Sarcocystis muriviperae''
* ''
Sarcocystis neotomafelis''
* ''
Sarcocystis nesbitti''
* ''
Sarcocystis neurona''
* ''
Sarcocystis odoi''
* ''
Sarcocystis odocoileocanis''
* ''
Sarcocystis ovicanis''
* ''
Sarcocystis ovifelis''
* ''
Sarcocystis oviformis''
* ''
Sarcocystis ovalis''
* ''
Sarcocystis phacochoeri''
* ''
Sarcocystis phoeniconaii''
* ''
Sarcocystis pilosa''
* ''
Sarcocystis podarcicolubris''
* ''
Sarcocystis poephagi''
* ''
Sarcocystis poephagicanis''
* ''
Sarcocystis porcifelis''
* ''
Sarcocystis porcihominis''
* ''
Sarcocystis rangi''
* ''
Sarcocystis rangiferi''
* ''
Sarcocystis rauschorum''
* ''
Sarcocystis rileyi''
* ''
Sarcocystis rodentifelis''
* ''
Sarcocystis roudabushi''
* ''
Sarcocystis scandinavica''
* ''
Sarcocystis sebeki''
* ''
Sarcocystis sibirica''
* ''
Sarcocystis sigmodontis''
* ''
Sarcocystis silva''
* ''
Sarcocystis singaporensis''
* ''
Sarcocystis speeri''
* ''
Sarcocystis stehlinii''
* ''
Sarcocystis stenodactylicolubris''
* ''
Sarcocystis strixi''
* ''
Sarcocystis suicanis''
* ''
Sarcocystis suihominis''
* ''
Sarcocystis sulawesiensis''
* ''
Sarcocystis sybillensis''
* ''
Sarcocystis tarandi''
* ''
Sarcocystis tarandivulpes''
* ''
Sarcocystis tenella''
* ''
Sarcocystis tilopodi''
* ''
Sarcocystis turcicii''
* ''
Sarcocystis turdi''
* ''
Sarcocystis ursusi''
* ''
Sarcocystis venatoria''
* ''
Sarcocystis villivilliso''
* ''
Sarcocystis wapiti''
* ''
Sarcocystis zamani''
Evolution
These protozoa are being increasingly well studied in mammals, birds and reptiles. They do not appear to infect mammals of the superorder
Afrotheria
Afrotheria ( from Latin ''Afro-'' "of Africa" + ''theria'' "wild beast") is a superorder of placental mammals, the living members of which belong to groups that are either currently living in Africa or of African origin: golden moles, elephan ...
and infect only two species of the
Xenarthra
Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a superorder and major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and ...
. Because of this pattern, the genus may have evolved in the Northern Hemisphere from a pre-existing protozoan species that infected mammals.
Alternatively, because a number of Australian
marsupials
Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a ...
are also infected by this genus, marsupials may have been the original hosts of this genus and the parasites were spread to the Northern Hemisphere by birds. A third possibility is that the genus originally infected birds and was spread worldwide by these hosts. A final possibility because of the existence of lifecycles where both the intermediate and final hosts are reptiles, the genus may have originated in reptiles and spread from there to other genera. The resolution of this question awaits the outcome of further molecular studies.
Clinical: Human
Infection with ''Sarcocystis'' is known as sarcosporidiosis or sarcocystosis. Because of initial confusion over the nature of this parasite, the organism in the intestine was originally referred to as ''Isospora hominis'' .
Epidemiology
Although human intestinal infection is common, extraintestinal human sarcocystosis is considered to be rare.
The extremes of age reported to date are a 26-day-old infant and a 75-year-old man. Infections have been reported from
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 ...
,
Europe
Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
(
Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Spain
Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
and
Poland
Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
), the
United States
The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
(
California
California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
), Central and
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
,
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 ...
,
India
India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Tibet
Tibet (; ''Böd''; ), or Greater Tibet, is a region in the western part of East Asia, covering much of the Tibetan Plateau and spanning about . It is the homeland of the Tibetan people. Also resident on the plateau are other ethnic groups s ...
,
Malaysia
Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
, and
Southeast Asia
Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
.
Stool examinations in Thai laborers showed that ''Sarcocystis'' infection had a high prevalence of around 23%, reflecting ingestion of raw or undercooked meat. Virtually all cases appeared to be asymptomatic. A study of 100 human tongues obtained ''post mortem'' in
Malaya revealed an infection rate of 21%. No sex difference was found and the age range was 16 to 57 years (
mean
A mean is a quantity representing the "center" of a collection of numbers and is intermediate to the extreme values of the set of numbers. There are several kinds of means (or "measures of central tendency") in mathematics, especially in statist ...
37.7 years).
A non-enteric outbreak affecting 93 people was reported in 2012 in Malaysia. ''
Sarcocystis nesbitti'' was confirmed to be the cause in several cases.
History
The first report of human infection was made in 1868. Although several additional reports were subsequently published, these early descriptions were not considered definitive. The first generally agreed definitive description of this disease was published in 1894 by Baraban and Saint-Remy.
[Baraban M. Le and Saint-Remy M. G.(1894) Sur un cas de tubes psorospermiques observés chez l'homme. Compt. Rend. Soc. Biol. 46: 231–203] This species was named by Rivolta after Lindemannl in 1898.
The cysts in human muscle were considered to belong to a single species, ''S. lindemanni'', and the intestinal form to be ''S. hominis'' (from undercooked beef) or ''S. suihominis'' (from undercooked pork). The
description
Description is any type of communication that aims to make vivid a place, object, person, group, or other physical entity. It is one of four rhetorical modes (also known as ''modes of discourse''), along with exposition, argumentation, and narr ...
of ''S. lindemanni'' has since been considered to be unsatisfactory and has been declared a ''
nomen nudum
In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'' (a name lacking a description). Two species are currently considered to be capable of causing human intestinal infection: ''S. bovihominis'' (''S. hominis'') and ''S. suihominis''. Other species including ''S. nesbitti'' have occasionally been reported as infecting humans extraintestinally.
Events occurring within animals
Between the years 1992 and 1996, an investigation was conducted where 605 sheep, 826 goats, 1080 cattle, 580 water buffaloes and 36 camels had been slaughtered. The event happened in the Baghdad area in Iraq, the research used naked eye examination for macroscopic Sarcocysts, and peptic digestion, muscle squash, squeezing methods and indirect fluorescent antibody tests for microscopic types. Dogs had also been used in the trials as well when they had consumed cysts containing the infectious parasite or any other microscopic lifeform.
The prevalence of these parasitic lifeforms including Sarcocysts(Sarcocystis) was as follows from the results, the macroscopic cysts were 4.1, 33.6, 0.2, 15.6 and 0, and of the microscopic type, 97.0, 97.4, 97.8, 82.9 and 91.6 for the above-mentioned hosts. The most effective method was peptic digestion, giving a 93.3% rate,
fluorescent antibody test (IFAT) (88.6%), squeezing (81.3%), and muscle squash (81.2%). And as for the dogs, each one had shed about a total of 150-120 million
sporocysts.
Route of infection
Intestinal infection occurs when raw or undercooked
meat
Meat is animal Tissue (biology), tissue, often muscle, that is eaten as food. Humans have hunted and farmed other animals for meat since prehistory. The Neolithic Revolution allowed the domestication of vertebrates, including chickens, sheep, ...
is ingested. Contaminated water might be a source of very rare human extraintestinal infection (it is not possible for water to be the origin of a gut infection), but this remains a theoretical possibility. Fecal matter is also another way of transportation from one host to another for the parasitic lifeform.
Pathology
The pathology is of two types: a rare invasive form with
vasculitis
Vasculitis is a group of disorders that destroy blood vessels by inflammation. Both artery, arteries and veins are affected. Lymphangitis (inflammation of lymphatic vessels) is sometimes considered a type of vasculitis. Vasculitis is primarily c ...
and
myositis and an intestinal form that presents with
nausea
Nausea is a diffuse sensation of unease and discomfort, sometimes perceived as an urge to vomit. It can be a debilitating symptom if prolonged and has been described as placing discomfort on the chest, abdomen, or back of the throat.
Over 30 d ...
, abdominal pain, and
diarrhea
Diarrhea (American English), also spelled diarrhoea or diarrhœa (British English), is the condition of having at least three loose, liquid, or watery bowel movements in a day. It often lasts for a few days and can result in dehydration d ...
. While normally mild and lasting under 48 hours, the intestinal form may occasionally be severe or even life-threatening. The invasive form may involve a wide variety of tissues including lymph nodes, muscles, and the
larynx
The larynx (), commonly called the voice box, is an organ (anatomy), organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal ...
.
Clinical features
In volunteer studies with infected beef, symptoms appeared 3–6 hours after eating. These included anorexia, nausea, abdominal pain, distension, diarrhea, vomiting, dyspnea, and tachycardia. All symptoms were transient and lasted about 36 hours. In a second series, symptoms—abdominal pain, distension, watery diarrhea, and eosinophilia—appeared at 1 week and resolved after 3 weeks.
Clinical cases have been associated with acute fever, myalgias, bronchospasm, pruritic rashes, lymphadenopathy, subcutaneous nodules associated with eosinophilia, elevated erythrocyte sedimentation rate, and elevated creatinine kinase levels. Symptoms may last as long as five years. Segmental necrotizing enteritis has been reported on one occasion.
Diagnosis
Definitive diagnosis by
biopsy
A biopsy is a medical test commonly performed by a surgeon, interventional radiologist, an interventional radiologist, or an interventional cardiology, interventional cardiologist. The process involves the extraction of sampling (medicine), sample ...
of an infected muscle. Sarcocysts are identifiable with
hematoxylin
Haematoxylin American and British English spelling differences#ae and oe, or hematoxylin (), also called natural black 1 or Colour Index International, C.I. 75290, is a chemical compound, compound extracted from wood#Heartwood and sapwood, heart ...
and
eosin
Eosin is the name of several fluorescent acidic compounds which bind to and from salts with basic, or eosinophilic, compounds like proteins containing basic amino acid residues such as histidine, arginine and lysine, and stains them dark red ...
. The PAS stain may be helpful, but variable uptake of stain is common. Along with the sarcocysts, inflammatory cells may be found. Other findings include myositis, myonecrosis, perivascular and interstitial inflammation, vasculitis, and eosinophilic myositis.
Prevention
Infection can be prevented by cooking the meat before eating. Alternatively, freezing the meat at −5 °C for several days before ingestion kills the sporocysts.
Clinical: Nonhuman

Four recognised species infect cattle: ''S. bovifelis'', ''S. bovihominis'' (''S. hominis''), ''S. cruzi'' (''S. bovicanis''), and ''S. hirsuta''. ''S. cruzi'' is the only species known to be pathogenic in cattle. Several clinical syndromes have been reported in connection with this parasite: eosinophilic myositis;
abortion
Abortion is the early termination of a pregnancy by removal or expulsion of an embryo or fetus. Abortions that occur without intervention are known as miscarriages or "spontaneous abortions", and occur in roughly 30–40% of all pregnan ...
s, stillbirths, and deaths in pregnant cows; two cases of necrotic
encephalitis
Encephalitis is inflammation of the Human brain, brain. The severity can be variable with symptoms including reduction or alteration in consciousness, aphasia, headache, fever, confusion, a stiff neck, and vomiting. Complications may include se ...
in heifers have also been reported. Typical clinical signs of acute bovine sarcocystosis are:
anorexia
Anorexia nervosa (AN), often referred to simply as anorexia, is an eating disorder characterized by Calorie restriction, food restriction, body image disturbance, fear of gaining weight, and an overpowering desire to be thin.
Individuals wit ...
,
pyrexia (42 °C or more),
anemia
Anemia (also spelt anaemia in British English) is a blood disorder in which the blood has a reduced ability to carry oxygen. This can be due to a lower than normal number of red blood cells, a reduction in the amount of hemoglobin availabl ...
,
cachexia
Cachexia () is a syndrome that happens when people have certain illnesses, causing muscle loss that cannot be fully reversed with improved nutrition. It is most common in diseases like cancer, Heart failure, congestive heart failure, chronic o ...
, enlarged palpable
lymph node
A lymph node, or lymph gland, is a kidney-shaped organ of the lymphatic system and the adaptive immune system. A large number of lymph nodes are linked throughout the body by the lymphatic vessels. They are major sites of lymphocytes that includ ...
s, excessive salivation, and loss of hair at the tip of the tail.
Sheep may be infected by four recognized species of ''Sarcocystis'': ''S. arieticanis'' and ''S. tenella'' (''S. ovicanis'') are pathogenic; ''S. gigantea'' (''S. ovifelis'') and ''S. medusiformis'' are nonpathogenic. Infection with these parasites is common in the US with over 80% of sheep examined showing evidence of infection.
''S. arieticanis'' and ''S. tenella'' both produce extraintestinal disease. Anemia, anorexia, ataxia, and abortions are the chief clinical signs. Myositis with flaccid
paralysis
Paralysis (: paralyses; also known as plegia) is a loss of Motor skill, motor function in one or more Skeletal muscle, muscles. Paralysis can also be accompanied by a loss of feeling (sensory loss) in the affected area if there is sensory d ...
has been reported as a consequence of infection.
Ovine protozoan myeloencephalitis is a recognised syndrome that may occur in outbreaks. The usual pathological findings in such cases are multifocal
spinal cord
The spinal cord is a long, thin, tubular structure made up of nervous tissue that extends from the medulla oblongata in the lower brainstem to the lumbar region of the vertebral column (backbone) of vertebrate animals. The center of the spinal c ...
white matter oedema and necrosis, glial nodules and mild to moderate nonsuppurative
encephalomyelitis
Encephalomyelitis is inflammation of the brain and spinal cord. Various types of encephalomyelitis include:
* '' Acute disseminated encephalomyelitis'' or ''postinfectious encephalomyelitis'', a demyelinating disease of the brain and spinal cord ...
. The diagnosis may be established finding protozoan bodies (12.7–23.0 μm) that stain immunocytochemically for ''Sarcocystis'' epitopes.
Four recognised species infect pigs: ''S. medusiformis'', ''S. meischeriana'' (''S. suicanis''), ''S. porcifelis'', and ''S. suihominis''. ''S. porcifelis'' is pathogenic for pigs causing diarrhea, myositis and lameness.
Five species infect horses: ''S. asinus'', ''S. bertrami'', ''S. equicanis'', ''S. fayeri'', and ''S. neurona'' (''S. falcatula''). All use canids as definitive hosts; transplacental infection has also been reported. ''S. neurona'' causes
equine protozoal myeloencephalitis. Exposure to this parasite appears to be common in the United States, with
serological surveys indicating that 50–60% percent of all horses in the Midwest United States have been exposed to it. Clinical signs include gait abnormalities including ataxia, knuckling, and crossing over. Muscle atrophy, usually unilateral, may occur. The lesions are typically focal. Brain stem involvement is common. Depression, weakness, head tilt, and dysphagia also occur.
''S. fayeri'' may cause myositis in horses.
Fatal infection of an alpaca (''
Lama pacos'') with an unnamed species has been reported. Findings included disseminated eosinophilic myositis, abortion, and haemoabdomen. The myositis was associated with haemorrhage, necrosis, and degeneration.
Infection by ''S. tilopodi'' of muscle tissue in the
guanaco
The guanaco ( ; ''Lama guanicoe'') is a camelid native to South America, closely related to the llama. Guanacos are one of two wild South American camelids; the other species is the vicuña, which lives at higher elevations.
Etymology
The gua ...
has been reported.
''S. hemionilatrantis'' infects mule deer. Death from experimental inoculation has been reported.
These parasites can also infect birds, producing three different clinical forms: an acute pulmonary disease, muscular disease, and neurological disease. Symptoms include lethargy, shortness of breath, tail bobbing, yellow-tinted droppings, and sudden death. The presence of the cysts in the muscle of wild birds is known as "rice breast".
Incidence in animals

Infection with ''Sarcocystis'' is common. Rates in pigs vary: 18% in
Iowa
Iowa ( ) is a U.S. state, state in the upper Midwestern United States, Midwestern region of the United States. It borders the Mississippi River to the east and the Missouri River and Big Sioux River to the west; Wisconsin to the northeast, Ill ...
,
27% in the
Philippines
The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
,
43% in Spain,
57% in
Uruguay
Uruguay, officially the Oriental Republic of Uruguay, is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast, while bordering the Río de la Plata to the south and the A ...
,
and 68% in India
The infection rate in sheep is commonly above 90%.
Camels have a similarly high incidence of infection.
Rates above 80% are known in cattle and goats.
The incidence in water buffaloes, yak and
hainag exceeds 80%
while the incidence in horses, donkeys, and chickens is lower.
Diagnosis
The diagnosis is usually made ''post mortem'' by examination of the skeletal muscle. In some species, the cysts may be visible to the naked eye (
duck
Duck is the common name for numerous species of waterfowl in the family (biology), family Anatidae. Ducks are generally smaller and shorter-necked than swans and goose, geese, which are members of the same family. Divided among several subfam ...
s,
mice
A mouse (: mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
,
rabbit
Rabbits are small mammals in the family Leporidae (which also includes the hares), which is in the order Lagomorpha (which also includes pikas). They are familiar throughout the world as a small herbivore, a prey animal, a domesticated ...
s and
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 ...
), but in most, microscopic examination is required.
''Ante mortem'' diagnosis may be made with the use of dermal sensitivity testing or
complement fixation tests. Muscle biopsy is also diagnostic, but this is much less commonly used.
Microscopy
Oocysts with two sporocysts or individual sporocysts in human feces are diagnostic of intestinal infection. These first appear 14 to 18 days after ingesting beef (''S. hominis''), and 11 to 13 days after ingesting pork (''S. suihominis''). Flotation based on high-density solutions incorporating sodium chloride, cesium chloride, zinc sulfate, sucrose, Percoll, Ficoll-Hypaque, or other such density gradient media is preferred to formalin-ethyl acetate or other sedimentation methods. Sporocysts of ''S. hominis'' average 9.3 by 14.7 μm and those of ''S. suihominis'' average 10.5 by 13.5 μm. Because of the overlap in size, size alone is not reliable as a diagnostic criterion of the species. Confirmatory staining with the periodic acid-Schiff (PAS) can be performed, as the walls stain positively. Heated safranin + methylene blue has been used for staining, as well.
PCR amplification of the
rRNA
Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
may also be used.
Cell walls
The walls of the sarcocyst may be helpful in species diagnosis with 24 wall types identified in 62 species. ''S. hominis'' and ''S. suihominis'' both have walls of type 10. The wall of ''S. hominis'' is up to 6 μm thick and appears radially striated from villar protrusions up to 7 μm long. Its bradyzoites are 7 to 9 μm long. The wall of ''S. suihominis'' is 4 to 9μm thick, with villar protrusions up to 13 μm long. Its bradyzoites are 15 μm long.
Differential diagnosis
Several other genera of heteroxenous and cyst-forming coccidia are known, including ''
Besnoitia'', ''
Cystoisospora'', ''
Frenkelia'', ''
Hammondia'', ''
Neospora'' and ''
Toxoplasma''. Related but
monoxenous spore-forming genera include ''Isospora''. Differentiating these genera from ''Sarcocystis'' in diagnostic material may be difficult without immunochemical stains.
Treatment
Current treatments are not entirely satisfactory.
Amprolium
Amprolium is the organic compound sold as a coccidiostat used in poultry. It has many International Nonproprietary Names.
__TOC__
Uses in coccidiosis treatment in poultry
The drug is a thiamine analogue and blocks the thiamine transporter of '' ...
(100 mg/kg, daily for 30 days), fed
prophylactically, reduced illness in cattle inoculated with ''S. cruzi''.
Prophylactic
Preventive healthcare, or prophylaxis, is the application of healthcare measures to prevent diseases.Hugh R. Leavell and E. Gurney Clark as "the science and art of preventing disease, prolonging life, and promoting physical and mental health a ...
administration of
amprolium
Amprolium is the organic compound sold as a coccidiostat used in poultry. It has many International Nonproprietary Names.
__TOC__
Uses in coccidiosis treatment in poultry
The drug is a thiamine analogue and blocks the thiamine transporter of '' ...
or
salinomycin also protected experimentally infected sheep.
In horses, treatment has been confined to dihydrofolate reductase inhibitors such as the
sulfonamide
In organic chemistry, the sulfonamide functional group (also spelled sulphonamide) is an organosulfur group with the Chemical structure, structure . It consists of a sulfonyl group () connected to an amine group (). Relatively speaking this gro ...
s and
pyrimethamine
Pyrimethamine, sold under the brand name Daraprim among others, is a medication used with leucovorin (leucovorin is used to decrease side effects of pyrimethamine; it does not have intrinsic anti-parasitic activity) to treat the parasitic disea ...
. Sulfadiazine (20 mg/kg orally) once or twice a day is a commonly used. Infected horses should also be placed on pyrimethamine at the dose of 1.0 mg/kg given once a day orally for 120 days or longer.
Diclazuril and
toltrazuril
Toltrazuril is an antiparasitic medication used primarily to treat coccidiosis in animals. Coccidiosis is a parasitic disease caused by coccidia, which are microscopic, spore-forming, single-celled obligate intracellular parasites belonging to ...
and other coccidiostats are being evaluated to treat EPM.
Since infection is rarely symptomatic, treatment is rarely required. No trials have been published, so treatment remains empirical.
Amprolium
Amprolium is the organic compound sold as a coccidiostat used in poultry. It has many International Nonproprietary Names.
__TOC__
Uses in coccidiosis treatment in poultry
The drug is a thiamine analogue and blocks the thiamine transporter of '' ...
and
salinomycin were effective in preventing severe illness and death in experimentally infected calves and lambs. These agents have not been tried in humans to date.

Muscle Sarcocystis is a parasitic infection causing muscle inflammation, pain, and weakness. Diagnosis is difficult and may require a biopsy.
Albendazole
Albendazole is a broad-spectrum antihelmintic and antiprotozoal agent of the benzimidazole type. It is used for the treatment of a variety of intestinal parasite infections, including ascariasis, pinworm infection, hookworm infection, trichuri ...
has been effective in some cases.
Albendazole
Albendazole is a broad-spectrum antihelmintic and antiprotozoal agent of the benzimidazole type. It is used for the treatment of a variety of intestinal parasite infections, including ascariasis, pinworm infection, hookworm infection, trichuri ...
is a medication used to treat parasitic infections by disrupting the parasites' ability to absorb nutrients, leading to their elimination. It is effective against infections caused by worms and some protozoa, including conditions like tapeworms, roundworms, and muscle Sarcocystis.
While
Albendazole
Albendazole is a broad-spectrum antihelmintic and antiprotozoal agent of the benzimidazole type. It is used for the treatment of a variety of intestinal parasite infections, including ascariasis, pinworm infection, hookworm infection, trichuri ...
has been used to deter Sarcocystis, it is not a definitive cure, often used with combinations of other medicines such as
prednisone
Prednisone is a glucocorticoid medication mostly used to immunosuppressive drug, suppress the immune system and decrease inflammation in conditions such as asthma, COPD, and rheumatologic diseases. It is also used to treat high blood calcium ...
,
cotrimoxazole, and
methotrexate
Methotrexate, formerly known as amethopterin, is a chemotherapy agent and immunosuppressive drug, immune-system suppressant. It is used to treat cancer, autoimmune diseases, and ectopic pregnancy, ectopic pregnancies. Types of cancers it is u ...
.
Vaccination
No vaccines are currently known. Experimentally inoculated pigs appear to develop a persistent immunity, so a vaccine may be possible.
Host-parasite relations
The parasite's lifecycle typically involves a predator and a prey animal. A single species may infect multiple prey or predator animals. In at least 56 species, definitive and intermediate hosts are known. Many species are named after their recognised hosts.
A listing of the known host-parasite relations can be found on the page
Sarcocystis: Host-parasite relations.
Notes
''Hoareosporidium'' is now considered a synonym of ''Sarcocystis''. The original type species was ''Sarcocystis miescheriana''. Its description has since been considered less than satisfactory and ''S. muris'' has been proposed as the type species. ''S. turdi'' may not be a valid species.
''Isospora bigemina'' has been reclassified as ''Sarcocystis bigemina''. ''Isospora hominis'' has been reclassified as ''Sarcocystis hominis''. ''S. bovihominis'' is a synonym of ''S. hominis''. ''S. cruzi'' is a synonym of ''S. bovicanis''. ''S. gigantea'' is a synonym of ''S. ovifelis''. ''S. hirsuta'' is a synonym of ''S. bovifelis''. ''S. idahoensis'' and ''S. roudabushi'' may be the same species. ''S. miescheriana'' is a synonym of ''S. suicanis''. ''S. neurona'' is a junior synonym of ''S. falcatula''. ''S. neuroma'' is the more commonly used name for this species. ''S. poephagi'' may be the same species as ''S. hirsuta''. ''S. poephagicanis'' may be the same species as ''S. cruzi''. ''S. tenella'' is a synonym of ''S. ovicanis''.
See also
*
Equine protozoal myeloencephalitis
References
External links
*
''Sarcocystis'' genome project* Review:
{{Authority control
Apicomplexa genera
Conoidasida
Rodent-carried diseases
Parasitic diseases
Parasites of equines
Parasites of mammals
Parasitic diseases associated with beef and pork consumption