Pyrenocine A
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''Curvularia inaequalis'' is a plant
saprobe 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 ...
that resides in temperate and subtropical environments. It is commonly found in the
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
s of
forage grass Fodder (), also called provender (), is any agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food given to the animals (includin ...
es and
grain A grain is a small, hard, dry fruit (caryopsis) – with or without an attached husk, hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and ...
s. The species has been observed in a broad distribution of countries including Turkey, France, Canada, The United States, Japan and India. This species is
dematiaceous 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 a
hyphomycete Hyphomycetes are a form classification of fungi, part of what has often been referred to as fungi imperfecti, Deuteromycota, or anamorphic fungi. Hyphomycetes lack closed fruit bodies, and are often referred to as moulds (or molds). Most hyphomy ...
.


History and taxonomy

The ''Curvularia'' genus can be identified by its spiral borne phaeophragmospores, which contain both
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
end cells and disproportionately large cells. They possess
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 differing curvature and number of septa. ''C. inaequalis'' was first described in 1907 by ecologist
Cornelius Lott Shear Cornelius Lott Shear (March 26, 1865 February 2, 1956) was an American mycology, mycologist and plant pathologist who served as a senior pathologist at the USDA Bureau of Plant Industry. Born in Coeyman's Hollow, Albany County, New York, on Mar ...
. The fungus was isolated from diseased New Jersey cranberry pulp and termed ''Helminthosporium inaequale''. Later, during Karl Boedijin's taxonomic organization and grouping 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 ...
, he recognized a similarity between them and ''H. inaequale''. He recognized a morphological similarity between its
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 ...
and those of the ''lunata'' group within ''Curvularia'', and so renamed it ''C. inaequalis''. Recognition of the three-
septate In biology, a septum (Latin for ''something that encloses''; septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatrial se ...
curved conidia motivated the introduction of the now popularized name.


Growth and morphology

The species' spore producing cells take on a model of sympodial growth. Conidia grow through successive apices which end in a terminal prospore. Growth can be affected by static magnetic fields with field flux densities. Under these conditions, the number of
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 ...
are able to increase by a minimum of 68 percent. ''Curvularia inaequalis'' is a filamentous fungus, with 3 to 12 densely packed filaments. The species is mostly brown in appearance, with pale brown end cells. Conidia themselves, consist of 3-5 cells with thick cell walls and a larger central cell. The diameter of the conidia ranges from 10 to 30 micrometers and have a slight leading curvature. Overall the appearance of the species is described as looking "cottony" with clear branching cells. The species can be difficult to identify due to its similar appearance to both '' C.'' and '' geniculate''. Instead, sequencing of nuclear rRNA
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...
regions (ITS) can be done to achieve accurate identification.


Physiology

The optimal growth temperature for the species is 30°C. It is able to produce a multitude of chemical products with enzymatic properties. One
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 ...
produced is
chloroperoxidase Chloride peroxidase () is a family of enzymes that catalyzes the chlorination of organic compounds. This enzyme combines the inorganic substrates chloride and hydrogen peroxide to produce the equivalent of Cl+, which replaces a proton in hydrocar ...
, which can catalyze
halogenation In chemistry, halogenation is a chemical reaction which introduces one or more halogens into a chemical compound. Halide-containing compounds are pervasive, making this type of transformation important, e.g. in the production of polymers, drug ...
reactions. Chloroperoxidase secreted from ''C. inaequalis'' contains
vanadium Vanadium is a chemical element; it has Symbol (chemistry), symbol V and atomic number 23. It is a hard, silvery-grey, malleable transition metal. The elemental metal is rarely found in nature, but once isolated artificially, the formation of an ...
active site. The presence of the vanadium substrate
vanadate In chemistry, a vanadate is an anionic coordination complex of vanadium. Often vanadate refers to oxoanions of vanadium, most of which exist in its highest oxidation state of +5. The complexes and are referred to as hexacyanovanadate(III) and no ...
is essential for the function of chloroperoxidase. The compound
glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
however, acts as an
inhibitor Inhibitor or inhibition may refer to: Biology * Enzyme inhibitor, a substance that binds to an enzyme and decreases the enzyme's activity * Reuptake inhibitor, a substance that increases neurotransmission by blocking the reuptake of a neurotransmi ...
for both enzyme function and production. In its active form, the enzyme is able to then produce
hypochlorous acid Hypochlorous acid is an inorganic compound with the chemical formula , also written as HClO, HOCl, or ClHO. Its structure is . It is an acid that forms when chlorine dissolves in water, and itself partially dissociates, forming a hypochlorite an ...
, a strong
oxidizing agent An oxidizing agent (also known as an oxidant, oxidizer, electron recipient, or electron acceptor) is a substance in a redox chemical reaction that gains or " accepts"/"receives" an electron from a (called the , , or ''electron donor''). In ot ...
. It has been theorized that ''C. inaequalis'' utilizes chloroperoxidase and hypochlorous acid in combination to penetrate the host's cell wall. Other significant compounds produced include of B-galactosidase, and . The species is able to produce large amounts of
β-galactosidase β-Galactosidase (EC 3.2.1.23, beta-gal or β-gal; systematic name β-D-galactoside galactohydrolase) is a glycoside hydrolase enzyme that catalyzes hydrolysis of terminal non-reducing β-D-galactose residues in β-D-galactosides. (This enzym ...
, which can hydrolyze lactose in acid whey. ''C. inaequalis'' also contains 4-hydroxyradianthin and Curvularone A compounds which have been identified as potential anti-tumor agents.


Pathology and toxicology


Plant pathology and toxicology

''Curvularia inaequalis'' is known to cause leaf spot, also known as
Leaf Blight Blight is a specific symptom affecting plants in response to infection by a pathogenic organism. Description Blight is a rapid and complete chlorosis, browning, then death of plant tissues such as leaves, branches, twigs, or floral organs. Ac ...
. Symptoms of infection by ''C. inaequalis'' include the combination of oval shaped dark brown patches and leaf tip dieback. The infection slowly spreads causes necrosis until it has covered the entirety of the leaf. It results in the thinning of grass vegetation such as
Zoysia ''Zoysia'' (;"Zoysia."
entry at CollinsDictionary.com. Retrieved 5 September 2021.
, -, ...
-, Bent-, Bermuda- and Buffalo- grasses. Blighting is believed to be caused by two ''C. inaequalis''
mycotoxins A mycotoxin (from the Greek μύκης , "fungus" and τοξικός , "poisonous") is a toxic secondary metabolite produced by fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' is usually r ...
, Pyrenocines and . Pyrenocines A is the more potent of the two, stunting growth and causing necrosis in vegetation. Both cause leaf tip die back in turf grass and leaf leakage of
electrolyte An electrolyte is a substance that conducts electricity through the movement of ions, but not through the movement of electrons. This includes most soluble Salt (chemistry), salts, acids, and Base (chemistry), bases, dissolved in a polar solven ...
s in Bermuda grass.


Human pathology

''Curvularia inaequalis'' is typically a rare human pathogen. There are however, recorded medical cases that mention infection by the species. One such case is of an
Eosinophilic Eosinophilic (Greek suffix '' -phil'', meaning ''eosin-loving'') describes the staining of tissues, cells, or organelles after they have been washed with eosin, a dye commonly used in histological staining. Eosin is an acidic dye for stainin ...
fungal
rhinosinusitis Rhinosinusitis is a simultaneous infection of the nasal mucosa (rhinitis) and an infection of the mucosa of the paranasal sinuses (sinusitis). A distinction is made between acute rhinosinusitis and chronic rhinosinusitis. Background Because sin ...
in an immunocompromised male.
Endoscopic An endoscopy is a procedure used in medicine to look inside the body. The endoscopy procedure uses an endoscope to examine the interior of a hollow organ or cavity of the body. Unlike many other medical imaging techniques, endoscopes are insert ...
sinus surgery was required to remove a large polyposis. ''C. inaequalis'' was found to have grown favorably in the eosinophilic mucus. Oral
itraconazole Itraconazole, sometimes abbreviated ITZ, is an antifungal medication used to treat a number of fungal infections. This includes aspergillosis, blastomycosis, coccidioidomycosis, histoplasmosis, and paracoccidioidomycosis. It may be given ...
and other
corticosteroids Corticosteroids are a class of steroid hormones that are produced in the adrenal cortex of vertebrates, as well as the synthetic analogues of these hormones. Two main classes of corticosteroids, glucocorticoids and mineralocorticoids, are invol ...
successfully were administered to prevent reinfection. Another case of ''C. inaequalis'' causing disease includes
peritonitis Peritonitis is inflammation of the localized or generalized peritoneum, the lining of the inner wall of the abdomen and covering of the abdominal organs. Symptoms may include severe pain, swelling of the abdomen, fever, or weight loss. One pa ...
in an elderly patient. It is suggested that contraction of the fungus occurs due to contact with soils. Furthermore, a case of recorded aerosolized ''C. inaequalis'' in one Canadian home supports airborne movement of spores as an important mode of transfer. While many cases of infection due to soil contact with the genus ''Curvularia'', connection with the specific species has not yet been confirmed. Further studies are required to determine its human pathogen potential.


References

{{Taxonbar , from = Q10463928 Pleosporaceae Fungi described in 1907 Fungus species