Aphanoascus Fulvescens
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''Aphanoascus fulvescens'' is a
mould A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. No ...
fungus that behaves as a keratinophilic saprotroph and belongs to the
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
. It is readily isolated from soil and dung containing keratin-rich tissues that have been separated from their animal hosts. This organism, distributed worldwide, is most commonly found in areas of temperate climate, in keeping with its optimal growth temperature of . While ''A. fulvescens'' is recognized as a
geophilic Geophilic means soil loving or preferring the soil. This term is usually used when referring to certain types of fungi or molds that live in the soil. Many of these organisms are usually recovered from the soil but occasionally infect humans and ...
fungal species, it is also a facultative opportunistic pathogen. Although it is not a dermatophyte, ''A. fulvescens'' has occasionally been shown to cause onychomycosis infections in humans. Its recognition in the laboratory is clinically important for correct diagnosis and treatment of human dermal infections.


Taxonomy

''Aphanoascus fulvescens'' has faced many discrepancies and challenges for proper
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
placement. The naming and classification of ''A. fulvescens'' has been subject to taxonomic confusion since the original discovery of the species.Cano J, Guarro J. (April 1990). “The genus ''Aphanoascus''”. ''Mycological Research'' 94: 355–377 The genus ''
Aphanoascus ''Aphanoascus'' is a genus of fungi in the family Onygenaceae. It was circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle ...
'', to which ''A. fulvescens'' belongs, was first described in 1890 by
Hugo Zukal Hugo Zukal (1845–1900) was an Austrian lichenologist and mycologist. Born in Troppau (now Opava in the Czech Republic), he graduated from high school there in 1859, and was afterwards employed as a botanist until 1864. From 1864 until 1872, h ...
after isolating it from alligator dung in Vienna.Rippon JW, Lee FC, McMillen S. (1970). “Dermatophyte infection caused by ''Aphanoascus fulvescens''”. ''Archives of Dermatology'' 102(5):552-&. doi:10.1001/archderm.102.5.552 Zukal named this fungus ''Aphanoascus cinnabarinus''. The same fungus however, had already been isolated earlier in 1875 under the name ''Badhamia fulvescens'' by Cooke. The genus ''
Aphanoascus ''Aphanoascus'' is a genus of fungi in the family Onygenaceae. It was circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle ...
'' was then further reviewed by Apinis in 1968 who deemed ''A. fulvescens'' as the type species. While most
mycologist Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, their Taxonomy (biology), taxonomy and ethnomycology, their use to humans, including as a so ...
s accept this title, others argue that ''Aphanoascus'' is a synonym for ''Anixiopsis'', which was the genus name given to a soil isolate described by Hansen as ''Anixiopsis fulvescens''. Later in 1973, further confusion was caused with the introduction of ''Aphanoascus cinnabarinus'' as the new type species for the genus ''Aphanoascus'' by Udagawa & Takada. In 1980, Benny & Kimbrough recognized ''Aphanoascus'' and ''Anixiopsis'' as two distinct genera, the former distinguished by reticulate ascospores and the latter by ridged ascospores. Whether the two genera are synonymous remains controversial; many authors continue to accept them as one genus. Initially, ''A. fulvescens'' was placed in the family
Cephalothecaceae The Cephalothecaceae are a family of fungi in the class Sordariomycetes. The family was circumscribed in 1917 by Austrian naturalist Franz Xaver Rudolf von Höhnel. Species in this family are saprobic, often growing on rotten wood or on other fun ...
by Apinis, in the Onygenaceae by Malloch & Cain and Currah, in Trichocomaceae by Benny & Kimbrough, and in Amauroascaceae by Arx. Today, most mycologists accept ''A. fulvescens'' as the type species of the genus ''Aphanoascus'' under the family Onygenaceae. ''A. fulvescens'' refers to the sexual state, or teleomorph, of the organism. ''
Chrysosporium keratinophilum ''Chrysosporium keratinophilum'' is a mold that is closely related to the dermatophytic fungi (Family Arthrodermataceae) and is mainly found in soil and the coats of wild animals to break down keratin. ''Chrysosporium keratinophilum'' is one o ...
'' is regarded by some mycologists as the asexual state, or
anamorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an ase ...
, however others argue that this is only the asexual state of ''Aphanoascus keratinophilus''.Campbell, C. K., Johnson, E. M. and Warnock, D. W. (2013) "Miscellaneous Moulds".''Identification of Pathogenic Fungi Second Edition''. Wiley-Blackwell. Oxford, UK. ''Aphanoascus fulvescens'', ''Aphanoascus terreus'', ''Aphanoascus canadensis'', ''Aphanoascus reticulisporus'', and ''Keratinophyton durum'' or ''Anixiopsis biplanata'' are the species that are currently accepted under the genus ''Aphanoascus''.


Appearance

The colonies of ''Aphanoascus fulvescens'' at on glucose-
peptone Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. A p ...
agar have a flat topology and a powdery to felt-like texture and are usually white to cream coloured depending on the number of brown ascomata present on the surface. Under the microscope, ''A. fulvescens'' is seen to have numerous, large club-shaped
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
with older colonies containing fruiting bodies, or ascomata, at their centres. These fruiting bodies are relatively large, ranging from 290–500 
μm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
in diameter, and are either colourless or have a light brown pigment. The presence of the ascomata is characteristic of all members of the
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
. The ascomata are smooth and have thick, enclosed walls lacking any openings. They reach full maturity after 8–10 days of colonial growth at optimal temperatures and eventually burst to release the
asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...
contained within them. The asci have walls composed of two to four layers of flattened cells which in total amount to 4–6 μm in thickness. There are eight ascospores contained within each
ascus An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or s ...
in ''A. fulvescens'' colonies. The
ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a surface that may be defined as the ...
al ascospores are a light brown colour and are either lens shaped or disc shaped. They can be as large as 5 x 3.5 μm and have notably rough walls. The ascospores of ''A. fulvescens'' are different than those of other members of the Amauroascaeae which have ascospores that are usually globular and round. Rather, the unicellular ascospores of ''A. fulvescens'' are found to be flattened with reticulate walls. The
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 or ...
of ''A. fulvescens'' are
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
, branched, and contain many cross-walls (setpa). They are generally 1.7–3 μm wide and are contained within thin walls. The colonial appearance of ''A. fulvescens'' generally mimics that of species in the genus ''
Trichophyton ''Trichophyton'' is a genus of fungi, which includes the parasitic varieties that cause tinea, including athlete's foot, ringworm, jock itch, and similar infections of the nail, beard, skin and scalp. Trichophyton fungi are molds characterized ...
'' and ''
Chrysosporium ''Chrysosporium'' is a genus of hyaline hyphomycetes fungi in the family Onygenaceae. ''Chrysosporium'' colonies are moderately fast-growing, flat, white to tan to beige in color; they often have a powdery or granular surface texture. Hyaline, ...
'' while under the microscope the appearance of the organism resembles that of the genus '' Aspergillus''. The differentiation of ''A. fulvescens'' from Aspergillus spp. is based on their
conidial A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to t ...
states.


Ecology

''Aphanoascus fulvescens'' is commonly found in the soil and in dung, living as a keratinophilic saprotroph. It is also often isolated from keratin-rich tissues such as hair and nails that have been discarded from the host. ''A. fulvescens'' is a
geophilic Geophilic means soil loving or preferring the soil. This term is usually used when referring to certain types of fungi or molds that live in the soil. Many of these organisms are usually recovered from the soil but occasionally infect humans and ...
fungus and therefore does not normally infect mammals. The organism is more commonly isolated from soil that has been inhabited frequently by animals than soil that has not. However, while this fungus is not a true dermatophyte, it is opportunistic and has been seen to cause dermatophytosis in humans and other mammals on occasion.Marin G, Campos R. (1984). “Dermatophytosis due to ''Aphanoascus fulvescens''”. ''Sabouraudia Journal of Medical and Veterinary Mycology'' 22(4):311–14. ''A. fulvescens'' has a worldwide distribution, however it is more frequently isolated in temperate climates. This is because the organism has optimal growth at 28 °C at which it grows 3–4 mm per day and up to 25 mm per week. At elevated temperatures, such as that of the human body (37 °C), ''A. fulvescens'' shows restricted growth, averaging only 5–6 mm of growth over the course of seven days. This may be a possible explanation as to why infections involving ''A. fulvescens'' are uncommon in humans and other mammals.


Pathogenicity

While ''Aphanoascus fulvescens'' is a
geophilic Geophilic means soil loving or preferring the soil. This term is usually used when referring to certain types of fungi or molds that live in the soil. Many of these organisms are usually recovered from the soil but occasionally infect humans and ...
fungus, it is also opportunistic and therefore has been isolated from humans and animals on occasion. In order to cause infection, the fungus must first adhere to the host cells. '' In vivo'' and '' in vitro'' experiments have proposed the possible role of secreted serine proteases in order to mediate this adhesion. Adhesion of ''A. fulvescens'' to the host cell is necessary, because the organism cannot degrade keratin from a distance and instead uses pressure to penetrate the keratin-rich cells of the host.Cano J, Guarro J, Mayayo J. (January 1990). “Experimental Pathogenicity of ''Aphanoascus'' spp.”. ''Mycoses'' 33(1): 41–45.


Skin

Keratinophilic fungal species that fall under the genus ''
Trichophyton ''Trichophyton'' is a genus of fungi, which includes the parasitic varieties that cause tinea, including athlete's foot, ringworm, jock itch, and similar infections of the nail, beard, skin and scalp. Trichophyton fungi are molds characterized ...
'', '' Epidermophyton'', or '' Microsporum'', are
pathogenic In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
to humans and other animals, and are referred to as dermatophytes on the basis of their ability to cause dermatophytosis in these hosts.de Vries G. (1962). “Keratinophilic fungi and their action”. ''Antonie van Leeuwenhoek'' 28:121–33 However, infections in these hosts caused by soil-derived keratinophilic saprotrophs like ''A. fulvescens'' are very seldom and only a few cases have been documented in the literature. The first documented case of ''A. fulvescens'' causing tinea corporis was in 1970 when the fungus was repeatedly isolated from a 4 x 5 cm red patch on the inner right thigh of a 21-year-old white male. This case showcases the clinical importance of non-dermatophyte keratinophilic fungi such as ''A. fulvescens'' as a result of the ability to cause dermatophytosis in the form of tinea corporis in humans. While visually, the case of tinea corporis was indistinguishable from those caused by true dermatophytes, the infection did not respond well to typical treatment. This observation is crucial for correct diagnosis and treatment of the skin lesion and highlights the necessity of awareness for atypical opportunistic human pathogenic fungi.


Hair

The
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
is the first region of the hair that is attacked in an ''A. fulvescens'' infection.English MP. “Destruction of hair by ''Chrysosporium keratinophilum''”. (April 1969). ''Transactions of the British Mycological Society'' 52(2):247–255.Cano J. Guarro J, Figueras M. (March 1991). "Study of the invasion of human hair in vitro by ''Aphanoascus'' spp." ''Mycoses''. 34(3):145–152. This attack resembles that of true dermatophytes in that some lateral
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 or ...
give rise to fronded mycelium. However, some
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 or ...
also give rise to
appressoria An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capable ...
, or flattened hyphal pressing organs, which are characteristic of non-keratinophilic fungi. Dermatophytes digest the hair cuticle rapidly, unlike ''A. fulvescens'', which breaks the cuticle down so slowly that the scales are still visible for weeks after the hair has been infected. After the cuticle has been broken down, the next structure that is targeted is the medulla. This region is first colonized by hyphae present in the cuticle and fungal growth is more rapid here because the
trichohyalin Trichohyalin is a protein that in mammals is encoded by the ''TCHH'' gene. Discovery In 1903 the name ''trichohyalin'' was assigned to the granules of the inner root sheath (IRS) of hair follicles discovered by Hans Vörner. In 1986 the name was ...
of the medulla is more readily digested than the keratin of the cuticle. The medullary mycelium then grow back towards the cuticle in a manner that makes the origin of the mycelium undetectable after 5–6 days of infection. The cortex is the third, and last region of the hair to be attacked by ''A. fulvescens''.
Appressoria An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capable ...
penetrate through to the cortex to produce cells of various sizes, giving the cortex a swollen morphology. This swollen mass of cells can proliferate in all directions to give rise to a large mass of mycelium that protrudes backwards out of the hair. Dermatophytes are seen to behave in a similar manner, however, instead of a bulging mass, the mycelium grows in a more organized fashion, as a single collapsed column. Growth of the cortical mycelium can be at any angle to the hair in an attack from ''A. fulvescens'', while true dermatophytes have the characteristic of growing along the hair in parallel. ''A. fulvescens'' can attack at any point along the hair and often causes a marked swelling upon entry sites. The mycelium penetrates the hair by exerting a high amount of pressure on the hair cells and is always closely pressed to the cavity walls. ''A. fulvescens'' cannot digest keratin from a distance, as seen in dermatophytes, but rather seems to require direct contact with the keratin-rich substrate. The digestion of keratin by ''A. fulvescens'' happens in two stages, the first of which involves the denaturation of keratin. The second stage involves the actual digestion of keratin by ''A. fulvescens'', which then grows into the remaining space. An infected hair is completely replaced by the mycelium of ''A. fulvescens'' within 6–8 weeks.


Treatment

Topical steroids and
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
ointments A topical medication is a medication that is applied to a particular place on or in the body. Most often topical medication means application to body surfaces such as the skin or mucous membranes to treat ailments via a large range of classes ...
such as Neosporin have not only shown to be ineffective in treating dermatophytosis caused by ''Aphanoascus fulvescens'', but have also been shown to enhance its fungal growth. Ringworm infections caused by ''A. fulvescens'' in animals have been successfully treated with
anti-fungal drug An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as crypto ...
s such as Tinactin, which contains tolnaftate as its active ingredient, and Grisactin, which contains griseofulvin as its active ingredient. However, topical treatment of human ''A. fulvescens'' infections with tolnaftate-containing drugs has been shown to worsen the lesions, causing further inflammation and increased sensitivity. Treatment with griseofulvin has proven somewhat effective in reducing lesion sizes and appears to have a dose-dependent effect. While lesions continue to heal after increased griseofulvin administration, there is some residual scarring of the infected area. Currently, there is no documentation providing evidence of an anti-fungal treatment that is completely effective against human dermatophytosis caused by ''A. fulvescens''.


References

{{Taxonbar, from=Q10414566 Fungi described in 1875 Onygenales