Blastocystis
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''Blastocystis'' is a genus of single-celled
heterokont Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which a ...
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
s belonging to a group of organisms that are known as the
Stramenopiles Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
(also called Heterokonts) that includes
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
,
diatom A diatom ( Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising se ...
s, and
water mold Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s. Blastocystis consists of several species, living in the
gastrointestinal tract 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 GI tract contains all the major organs of the digestive system, in humans and ...
s of species as diverse as humans, farm animals, birds, rodents, reptiles, amphibians, fish, and cockroaches. ''Blastocystis'' exhibits low host specificity, and many different species of ''Blastocystis'' can infect humans, and by current convention, any of these species would be identified as ''Blastocystis hominis''. ''Blastocystis'' is one of the most common human parasites in the world and has a global distribution. It is the most common parasitic infection in the United States, where it infected approximately 23% of the total population during year 2000. In less developed areas, infection rates as high as 100% have been observed. High rates of infection are found in individuals in developed countries who work with animals. Although the role of ''Blastocystis hominis'' in human disease is often referred to as controversial, a systematic survey of research studies conducted by 11 infectious disease specialists from nine countries, found that over 95% of papers published in the 10 years prior identified it as causing illness in immunocompetent individuals. The paper attributed confusion over pathogenicity to the existence of
asymptomatic carrier An asymptomatic carrier is a person or other organism that has become infected with a pathogen, but shows no signs or symptoms. Although unaffected by the pathogen, carriers can transmit it to others or develop symptoms in later stages of the d ...
s, a phenomenon the study noted is common to all gastrointestinal protozoa. However, ''Blastocystis'' has never fulfilled Koch's postulate that infection of a healthy individual with ''Blastocystis'' leads to disease. The fact that ''Blastocystis''' infection route is oral-anal indicates that carriers have been in contact with faecal contaminated matter which might have included other intestinal pathogens that explain the observed symptoms. A more likely explanation is the presence of virulent and non-virulent strains since there exists an enormous genetic variation between different strains (or genotypes). See the genotype paper by Rune Stensvold and the recent ''Blastocystis'' genome paper expanding on this diversity. An alternative theory that ''Blastocystis'' is not a pathogen at all has recently been strengthened based on its biochemistry.


Classification

The appropriate classification of ''Blastocystis'' has only recently been resolved. The original description of ''Blastocystis'' was as a
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
due to its yeast-like glistening appearance in fresh wet mounts and the absence of
pseudopod A pseudopod or pseudopodium (plural: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filament ...
ia and
locomotion Locomotion means the act or ability of something to transport or move itself from place to place. Locomotion may refer to: Motion * Motion (physics) * Robot locomotion, of man-made devices By environment * Aquatic locomotion * Flight * Locomo ...
. This was then contradicted by Zierdt, who reclassified it under
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of " subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists have also used th ...
Sporozoa The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. The ...
(and later in Sarcodina), based on some distinctive
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
an features of the ''Blastocystis'' cell. Its sensitivity to antiprotozoal drugs and its inability to grow on fungal media further indicated that it was a
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
n. However, major revisions were made to its classification. An analysis of gene sequences was performed in 1996, which placed it into the group
Stramenopiles Stramenopile is a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have be ...
. Other Stramenopiles include brown algae, mildew, diatoms, the organism that caused the Great Famine of Ireland, and the organism responsible for
Sudden oak death James Green aka "Sudden" is a fictional character created by an English author Oliver Strange in the early 1930s as the hero of a series, originally published by George Newnes Books Ltd, set in the American Wild West era. Oliver Strange died i ...
disease. However, the position of ''Blastocystis'' within the stramenopiles remains enigmatic.


Signs and symptoms

Most published studies have reported that between 50% and 80% of individuals mono-infected with ''Blastocystis'' will show symptoms._ Factors influencing presentation of symptoms have been listed as the patient's age, with younger patients less likely to show symptoms, as well as genetic changes that influence the production of cytokines. Some studies have suggested that pathogenicity may be linked to specific subtypes of ''Blastocystis'' and experimental infection of animals has reported varying degrees of illness depending on the subtype used. While some subtypes appear to be less likely to result in symptomatic infection, those subtypes are also found in symptomatic individuals who have no other infection found. Symptoms associated with the infection are diarrhea, constipation, nausea, abdominal cramps, bloating, excessive gas, and anal itching. Most cases of the infection appear to become diagnosed as
irritable bowel syndrome Irritable bowel syndrome (IBS) is a "disorder of gut-brain interaction" characterized by a group of symptoms that commonly include abdominal pain and or abdominal bloating and changes in the consistency of bowel movements. These symptoms may ...
, according to studies from Denmark, Pakistan, the United Kingdom, and Italy. The timescale of infection with the parasite can range from weeks to years. In the early 2000s, Egyptian physicians identified 84 patients with diarrhea and enteritis apparently caused by Blastocystis hominis. After three days of
nitazoxanide Nitazoxanide, sold under the brand name Alinia among others, is a broad-spectrum antiparasitic and broad-spectrum antiviral medication that is used in medicine for the treatment of various helminthic, protozoal, and viral infections. It is ind ...
treatment, symptoms cleared and no fecal organisms were detectable in 36 (86%) of 42 treated patients and in 16 (38%) of 42 people who received placebo (''P'' < .0001). The investigators concluded that either ''B hominis'' is pathogenic and can often be effectively treated with nitazoxanide, or that nitazoxanide (a drug approved by the FDA for the treatment of giardia and cryptosporidia) eradicated an unidentifiable organism.


Taxonomy

For many years, scientists believed one species of ''Blastocystis'' infected humans, while different species of ''Blastocystis'' infected other animals. So they called ''Blastocystis'' from humans ''Blastocystis hominis'' and gave different species names to ''Blastocystis'' from other animals, for example ''Blastocystis ratti'' from rats. In recent years, various genetic analysis have shown that ''Blastocystis hominis'' as a unique entity does not exist, i.e. there is no ''single'' species of ''Blastocystis'' that infects humans. In fact, a number of distinct genetic types of ''Blastocystis'' can infect humans, including those previously called ''Blastocystis ratti'' and the differences are so great that they could be considered separate species. Because of this, in 2007 scientists proposed discontinuing the use of the term ''Blastocystis hominis.'' Their proposal was to refer to ''Blastocystis'' from humans and animals as ''Blastocystis'' sp. subtype nn where nn is a number assigned to each group according to the degree of genetic identity of the ''Blastocystis'' organisms, based on gene sequences, rather than the host that was infected. At that time nine subtypes were known to infect mammals and birds, all of which had been found in humans. A tenth group was reported in China in 2007, but a full analysis of its relationships has not yet been performed and it is not yet clear whether it is a group within a described subtype or a new subtype. A definite tenth subtype was then found in a variety of other mammals, including primates, but it has not as yet been found in humans. There are now at least 13 genetically distinct small subunit ribosomal RNA lineages. These additional subtypes were found in a variety of mammalian hosts (including elephants and giraffes) and it is very likely that more subtypes will be found as more hosts are surveyed.


Epidemiology

''Blastocystis'' spp. prevalence in humans often exceeds 5% in industrialized countries. In the United States, it infected approximately 23% of the total population during year 2000. In less developed areas, infection rates with one or more subtypes are as high as 100%.


Transmission

Fecal-oral transmission is the most accepted pathway, and recent studies have shown that transmission involves only the cyst form of the parasite. The extent to which human-human, human-animal, and animal-human transmission occurs is still unknown. Genomic studies provide evidence for all three routes, though experimental studies have yet to provide conclusive proof for the existence of either.


Reservoir

Conclusively stating that ''Blastocystis'' has an animal reservoir depends upon unraveling the true nature of its transmission. If, as Noël et al. deem likely based upon their own molecular work and a review of the literature, animal-to-human transmission is possible, then animals such as pigs and dogs could in fact be acting as a large reservoir capable of human infection. Epidemiological studies finding that infection is more common in people living in proximity to farm animals or pets further supports this notion.


Morphology

''Blastocystis'' has various morphological forms. Four commonly described forms are the vacuolar (otherwise known as central body), granular, amoeboid, and cyst forms. The appearance of the organism is largely dependent upon environmental conditions as it is extremely sensitive to oxygen. Whether all of these forms exist in the host intestine is unclear. ;Vacuolar form The vacuolar form is the typical cell form of ''Blastocystis'' seen in culture and is often used for the identification of the organism. These vacuolar forms vary greatly in size, with
diameter In geometry, a diameter of a circle is any straight line segment that passes through the center of the circle and whose endpoints lie on the circle. It can also be defined as the longest chord of the circle. Both definitions are also valid f ...
s ranging between 2 µm and 200 µm. The vacuolar form is otherwise known as central body form because it has a large central
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic m ...
surrounded by a thin band of peripheral
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
which contains other organelles. Flocculent material has been described as being scattered unevenly throughout the vacuole. The function of the vacuole is still unclear, however, it has been suggested that, like for many
eukaryotic cells Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bact ...
, it is for storage purposes. Other functions, such as cell division during
reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual or ...
and the deposition of
apoptotic Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes ( morphology) and death. These changes incl ...
bodies, have been proposed, although more tests need to be done to validate these roles. ;Granular form The granular form is somewhere and somewhat morphologically similar to the vacuolar forms except that distinct granules are observed in the central vacuole and / or cytoplasm. Within the central vacuole, these granules appear in different forms too. Three types were suggested – metabolic,
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
, and reproductive granules. Metabolic granules play a role in chemical processes that are necessary for the maintenance of life in the organism. It was also put forward that reproductive granules were involved in the development of progeny cells. These hypotheses were made based on microscopy alone, which may be deemed misleading, hence more need to be done before making a definite conclusion. It has also been suggested that the granules may be an indication that the cell is dying. ;Amoeboid form The other form that exists is the amoeboid form. The amoeboid form of ''Blastocystis'' is non-motile and strongly adhesive. A research study has reported that amoeboid forms are produced only in cultures taken from symptomatic individuals, with asymptomatic individuals producing exclusively vacuolar forms. The study suggested this method could be used for diagnosing symptomatic infection. Additionally, it suggested the symptoms could be due to the accumulation of the strongly adhesive amoeboid forms on the host's intestinal wall. A detailed ultra-structural study of amoeboid forms was published in 2007. ;Cyst form The ''Blastocystis'' cyst form is a more recent discovery and has helped in the advancement of understanding the way the infection is transmitted. As compared to the other forms, it is generally smaller in size and has a thick multilayered cyst wall. It lacks a central vacuole and few nuclei, multiple vacuoles and food storage deposits were observed. The cyst form is the most resistant form of this parasite and is able to survive in harsh conditions because of its thick multilayered cyst wall. Experiments have been carried out to show its ability to withstand acidic gastric juices. Besides, the cysts did not lyse when placed in
distilled water Distilled water is water that has been boiled into vapor and condensed back into liquid in a separate container. Impurities in the original water that do not boil below or near the boiling point of water remain in the original container. Thus, di ...
and could survive well at
room temperature Colloquially, "room temperature" is a range of air temperatures that most people prefer for indoor settings. It feels comfortable to a person when they are wearing typical indoor clothing. Human comfort can extend beyond this range depending on ...
for up to 19 days, indicating its strong resistance.


Life cycle

The supposed life cycle begins with ingestion of the cyst form. After ingestion, the cyst develops into other forms which may in turn re-develop into cyst forms. Through human faeces, the cyst forms enter the external environment and are transmitted to humans and other animals via the faecal–oral route, repeating the entire cycle.


Obtaining and culturing ''Blastocystis''

The ATCC maintains a collection of ''Blastocystis'' isolates. Some records show whether the isolates were obtained from symptomatic or asymptomatic carriers. As yet, no publication has identified the subtypes of most of the ATCC isolates, which are mostly
axenic In biology, axenic (, ) describes the state of a culture in which only a single species, variety, or strain of organism is present and entirely free of all other contaminating organisms. The earliest axenic cultures were of bacteria or unicellu ...
. Researchers have reported that patients with ''Irritable bowel syndrome'' (IBS) may provide a reliable source for xenic ''Blastocystis'' isolates. Some researchers have reported being able to culture ''Blastocystis'' from 46% of IBS patients. Researchers have described different culture mechanisms for growing ''Blastocystis''. Colony growth on solid medium
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
on solid culture medium using a synthetic medium with added supplements have both been described. However, most cultivation is performed in liquid media of various types.


See also

* Blastocystosis * List of parasites (human)


References


External links


CDC description of ''Blastocystis hominis''

''Blastocystis'' Research Foundation

Dientamoeba Fragilis and Blastocystis Hominis

Blastocystis Blog by CR Stensvold
{{Taxonbar, from=Q1756496 Heterokont genera Parasitic SAR