Exophiala Pisciphila
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''Exophiala pisciphila'' is a
mesophilic A mesophile is an organism that grows best in moderate temperature, neither too hot nor too cold, with an optimum growth range from . The optimum growth temperature for these organisms is 37 °C (about 99 °F). The term is mainly applied ...
black yeast Black yeasts, sometimes also black fungi, dematiaceous fungi, microcolonial fungi or meristematic fungi is a diverse group of slow-growing microfungi which reproduce mostly asexually ( fungi imperfecti). Only few genera reproduce by budding cells ...
and member of the dark septate endophytes. This
saprotrophic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
fungus is found commonly in marine and soil environments. It is abundant in harsh environments like soil contaminated with heavy metals. ''E. pisciphila'' forms symbiotic relationships with various plants by colonizing on roots, conferring resistance to drought and heavy metal stress. It is an opportunistic pathogen that commonly causes infections in captive fish and amphibians, while rarely causing disease in
humans 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 ...
.
Secondary metabolites Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
produced by this species have potential clinical antibiotic and antiretroviral applications.


History and taxonomy

In 1969, Nikola Fijan first described a systemic
mycosis Fungal infection, also known as mycosis, is a disease caused by fungi. Different types are traditionally divided according to the part of the body affected: superficial, subcutaneous, and systemic. Superficial fungal infections include common ...
outbreak in
channel catfish The channel catfish (''Ictalurus punctatus''), known informally as the "channel cat", is a species of catfish native to North America. They are North America's most abundant catfish species, and the official state fish of Kansas, Missouri, Nebra ...
from a pond in
Alabama Alabama ( ) is a U.S. state, state in the Southeastern United States, Southeastern and Deep South, Deep Southern regions of the United States. It borders Tennessee to the north, Georgia (U.S. state), Georgia to the east, Florida and the Gu ...
and identified it as ''Exophiala salmonis''. In 1974, Michael McGinnis and Libero Ajello reevaluated the fungus and identified it as a new species ''Exophiala pisciphila''. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''pisciphila'' is a linguistic barbarism, combining the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word ''piscis'' meaning "fish" with the
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 ...
suffix ''-philos'' (φίλος) meaning "loving".


Habitat and ecology

''Exophiala pisciphila'' is commonly found in soil, plants and water in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
,
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
,
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Northwestern Europe, off the coast of European mainland, the continental mainland. It comprises England, Scotlan ...
, and
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. ''E. pisciphila'' occurs as a colonist or pathogen in cold-blooded vertebrates such as various commercially cultivated fish and amphibians. It has low host specificity. Captive fish are especially susceptible due to the confined space of aquariums and accumulation of fungal particles. Decorative pieces, stones or contaminated food in aquariums can all be reservoirs of ''E. pisciphila''. This fungus has a high tolerance to certain metals ions and has been encountered in harsh environments such as heavy metal polluted soils. When this fungus colonizes plant roots, it enhances plant tolerance to heavy metal ions.
Symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
relationships with host plants also allow for improved growth performance and plant survival rate in drought conditions.


Growth and morphology

''Exophiala pisciphila'' is an exclusively asexual fungus that exhibits both filamentous and yeast-like growth. Due to its variable growth forms and the dark pigmentation of its cell walls, it is considered a member of the descriptive grouping of similar fungi known as the
black yeast Black yeasts, sometimes also black fungi, dematiaceous fungi, microcolonial fungi or meristematic fungi is a diverse group of slow-growing microfungi which reproduce mostly asexually ( fungi imperfecti). Only few genera reproduce by budding cells ...
s. ''E. pisciphila'' forms slow growing colonies approximately in size which is similar to other species in the genus, ''E. salmonis'' and ''E. brunnea''. The texture of the colony is dry and fluffy due to the formation on aerial
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
in mature colonies. The upper surface is grey to green black in colour while the reverse surface tends to be black. Growth occurs on various media including malt extract agar (MA), oatmeal agar (OA), Sabourand's dextrose agar (SA), corn meal agar (CMA), Czapeck's solution agar,
potato dextrose agar Potato dextrose agar (BAM Media M127
from the ...
(PDA) and
nutrient agar Nutrient agar is a general-purpose solid medium supporting growth of a wide range of non- fastidious organisms. It typically contains ( mass/volume): * 0.5% peptone – this provides organic nitrogen * 0.3% beef extract/yeast extract – the ...
(NA). Optimal growth occurs on PDA and MA with the most aerial hyphae forming dome shaped colonies. Media interpreted to be associated with less optimal growth result in the formation of flat colonies. A distinguishing feature of this fungus from others in the genus is its ability to grow on L-arabinitol. Ideal growth conditions for ''E. pisciphila'' occur between , where maximum growth occurs at . This differentiates it from ''E. jeanselmei'' which has similar physiology otherwise. Reproduction for this species occurs asexually by conidiation which was observed to occur through various means in developing colonies. The conidia are produced either by (1) pre-existing conidia, (2) mature hyphae or (3) the differentiation of the cell into a specialized conidium-producing cell called an annellide. ''E. pisciphila'' have smooth-walled
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
with yellow-brown walls that characteristically differentiate into annelides. Annelides are bottle-shaped cells that give rise to conidia from a point at the tip of the bottle-neck, as it were. In this way, annelides are similar to
phialide The phialide ( ; , diminutive of phiale, a broad, flat vessel) is a flask-shaped projection from the vesicle (dilated part of the top of conidiophore) of certain fungi. It projects from the mycelium without increasing in length unless a subsequ ...
s but differ in that their necks incrementally elongate as each successive conidium is borne. The cell walls of this species contain the brown pigment
melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
which is both a pathogenicity factor and a mechanism of enhancing cell survival during periods of stress. The developing colonies also produce aerial hyphae that appear as hyphal strands that intertwine in a rope-like fashion. The formation of aerial hyphae has been suggested as a means of enhancing survival during harsh growth conditions. ''E. salmonis'' has single-celled conidia that are smaller than those of the otherwise morphologically the similar species, ''E. brunnea''.


Pathology

Unlike closely related species such as '' E. jeanselmei'' and '' E. dermatitidis'', ''E. pisciphila'' rarely causes disease in humans primarily due to its inability to tolerate human body temperature. One case of human disease was reported in
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 ...
where a person undergoing
immunosuppressive Immunosuppression is a reduction of the activation or efficacy of the immune system. Some portions of the immune system itself have immunosuppressive effects on other parts of the immune system, and immunosuppression may occur as an adverse react ...
therapy for a liver transplant developed a skin infection. The infection did not disseminate and resolved with therapy within a month. Uncontrolled
asthma Asthma is a common long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wh ...
tics may manifest
hypersensitivity Hypersensitivity (also called hypersensitivity reaction or intolerance) is an abnormal physiological condition in which there is an undesirable and adverse immune response to an antigen. It is an abnormality in the immune system that causes Imm ...
to ''E. pisciphila''
antigens In immunology, an antigen (Ag) is a molecule, moiety, foreign particulate matter, or an allergen, such as pollen, that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. An ...
. This fungus is pathogenic to an array of aquatic animals most notably freshwater and seawater fish in which infection is associated with the development of skin lesions and nodules on visceral organs. It can cause deadly infections in
Atlantic salmon The Atlantic salmon (''Salmo salar'') is a species of ray-finned fish in the family Salmonidae. It is the third largest of the Salmonidae, behind Hucho taimen, Siberian taimen and Pacific Chinook salmon, growing up to a meter in length. Atlan ...
where the hyphae invade the brain causing chronic inflammation. These infections are associated with abnormal swimming behaviours, depression and darkening of skin. Non-salmonid fish such as smooth dogfish,
channel catfish The channel catfish (''Ictalurus punctatus''), known informally as the "channel cat", is a species of catfish native to North America. They are North America's most abundant catfish species, and the official state fish of Kansas, Missouri, Nebra ...
,
American sole The American soles are a family (Achiridae) of flatfish occurring in both freshwater and marine environments of the Americas. The family includes about 35 species in seven genera. These are closely related to the soles (Soleidae), and have be ...
, Cardinal tetra,
cod Cod (: cod) is the common name for the demersal fish genus ''Gadus'', belonging to the family (biology), family Gadidae. Cod is also used as part of the common name for a number of other fish species, and one species that belongs to genus ''Gad ...
,
triggerfish Triggerfish are about 40 species of often brightly colored marine ray-finned fishes belonging to the family Balistidae. Often marked by lines and spots, they inhabit tropical and subtropical oceans throughout the world, with the greatest speci ...
, Japanese flounder, King George whiting,
American plaice The American plaice, American sole or long rough dab (''Hippoglossoides platessoides'') is a North Atlantic flatfish that belongs, along with other right-eyed flounders, to the family Pleuronectidae. In the northwest Atlantic (''H. p. platessoide ...
are also susceptible. Systemic, lethal infections have been described in captive sharks including the
zebra Zebras (, ) (subgenus ''Hippotigris'') are African equines with distinctive black-and-white striped coats. There are three living species: Grévy's zebra (''Equus grevyi''), the plains zebra (''E. quagga''), and the mountain zebra (''E. ...
,
bonnethead The bonnethead (''Sphyrna tiburo''), also called a bonnet shark or shovelhead, is a small member of the hammerhead shark genus ''Sphyrna'', and part of the family Sphyrnidae. It is an abundant species in the littoral zone of the North Atlantic a ...
and
hammerhead sharks The hammerhead sharks are a group of sharks that form the family Sphyrnidae, named for the unusual and distinctive form of their heads, which are flattened and laterally extended into a cephalofoil (a T-shape or "hammer"). The shark's eyes ar ...
. Infections of sharks, rays and skates are typically associated with severe tissue damage especially
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
of the spleen and gills. Other cold-blooded animals such as turtles, crabs, sea horses and frogs can be affected. ''E. pisciphila'' has been implicated as a minor egg pathogen due to its ability to infect a small number of
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
larvae. Isolates have been identified from tongue ulcers of various terrestrial animals such as horses and dogs.


Uses

''E. pisciphila'' produces Exophilin A, a
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
identified as a new
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting pathogenic bacteria, bacterial infections, and antibiotic medications are widely used in the therapy ...
against
Gram-positive bacteria In bacteriology, gram-positive bacteria are bacteria that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of cell wall. The Gram stain ...
. Another secondary metabolite produced by this species is a newly discovered
polyketide In organic chemistry, polyketides are a class of natural products derived from a Precursor (chemistry), precursor molecule consisting of a Polymer backbone, chain of alternating ketone (, or Carbonyl reduction, its reduced forms) and Methylene gro ...
compound 1-(3,5-dihydroxyphenyl)-4-hydroxypentan-2-one which may have antimicrobial activity. A novel fungal metabolite, Exophilic acid, has been isolated which acts as an inhibitor of HIV-1 integrase, an enzyme critical for replication and spread of HIV virus. This demonstrates its potential to be used for antiretroviral therapy.


References

{{Taxonbar, from=Q10493091 Chaetothyriales Fungi described in 1974 Fungus species