Zasmidium Cellare
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''Zasmidium cellare'', also known as cellar mold, is a species of fungus that exists in dark, ethanol-rich environments and is brown to black in colour. This species primarily exists in wine and brandy cellars in central and southern Europe, but can be found in surrounding regions and is thought to be helpful in the wine making process by some and a hygienic issue by others. Not much is known about ''Z. cellare'''s sexual biology and is thought to be beneficial to the cleanliness of cellar air due to its ability to consume musty odours.


Taxonomic history

''Z. cellare'' was originally classified by C. H. Persoon in 1794 under the name ''Racodium cellare'' – ''Racodium'' being a classification for plants that have no real relation or resemblance to each other. It would not be long, however, before Persoon retracted his original name, assigning ''
Antennaria ''Antennaria'' is a genus of dioecious perennial herbs in the family Asteraceae, native to temperate regions of the Northern Hemisphere, with a few species (''A. chilensis'', ''A. linearifolia'', ''A. sleumeri'') in temperate southern South ...
cellaris'' - another plant genus designation. It wasn't until Elias Fries that this species was designated correctly as a fungus under the name '' Zasmidium cellare'' in 1849. Over the years, there has been much contention over this classification, however. The first was Schanderl in 1936 who claimed that the genus '' Cladosporium'' was more fitting than '' Zasmidium''. Though ''Cladosporiums'' can be common indoor molds with brown or black colonies and have dark, pigmented
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 ...
, that is where the similarities with ''Z.'' ''cellare'' end. '' Cladosporiums'' usually exist on plant material and their
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s are often air dispersed, having a large abundance in outdoor environments, which simply isn't the case for ''Z. cellare.'' The second and third claim that ''Z. cellare'' was characterized incorrectly both occurred in 1971 by M. B. Ellis and Hawksworth who proposed ''Rhinocladiella'' ''cellaris'' and ''Rhinocladiella ellisii'', respectively. Hawksworth along with Riedl in 1977 re-proposed ''Rhinocladiella ellisii'', but in 1979 was criticized by De Hoog as the genus ''Rhinocladiella'' characterized ''Z. cellare's'' asexual (conidial) form of which the fungus rarely presents in and decided that ''Z. cellare'' was the most appropriate name for this species. To prevent any further contention, de Hoog amended the genus '' Zasmidium'' to include fungi with undifferentiated conidiogenous cells with wavy branches, ''"denticulate rachis",'' and pigmented scars. Today, the literature agrees that the proper classification is in fact ''Zasmidium cellare'' of the division ''
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 ...
'', representing spore shooting fungi; the class ''
Dothideomycetes Dothideomycetes is the largest and most diverse class of ascomycete fungi. It comprises 11 orders 90 families, 1300 genera and over 19,000 known species. Traditionally, most of its members were included in the loculoascomycetes, which is not par ...
'', which are fungi that grow in what are considered hostile or non-optimal conditions to most fungal species; the order '' Capnodiales'', which typically grow masses of black cells; and the family '' Mycosphaerellaceae'', which is a grouping of sac fungi.


Growth and morphology


Growth inhibition

''Z. cellare'' shares many morphological characteristics with another fungus, ''
Cladosporium sphaerospermum ''Cladosporium sphaerospermum'' is a radiotrophic fungus belonging to the genus ''Cladosporium'' and was described in 1886 by Albert Julius Otto Penzig from the decaying leaves and branches of ''Citrus''. It is a dematiaceous (darkly-pigmented) ...
'' (commonly found on shower ceilings and can live off of the oil in paints). Though ''C. sphaerospermum'' is very hard to get rid of with rigorous cleaning, ''Z. cellare'' requires very little in the way of interference to inhibit its growth, as even the slightest bit of care for the cleanliness of a cellar can prevent traces of the organism. The use of steel tanks for aging is also another inhibitor of ''Z. cellare'' growth, as steel is less porous than wood and prevents alcohol vapour from diffusing into the environment, and alone can be the soul contributor to the species' extinction as claimed by Henry Tribe (2006).


Morphology

''Z. cellare'' growth typically includes
hypha 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 ...
e (up to 5000 μm in length with walls 0.5-0.6 μm thick) with very small conidiophores, often not distinguishable from vegetative
hypha 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 ...
e, upon which
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s are produced at the end of teeth-like structures that are less pigmented than the basal portion of the conidiophore. Aerial hyphae are usually rough and darkly coloured, 2-2.5 μm wide, with thick walls, while the submerged hyphae are smooth and 2-3 μm wide. The Aerial hyphae are observed to have conidiogenous cells (thought to be remnants of conidiophores) primarily on the terminal of the
hypha 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 ...
e, sometimes on the sides, that are cylindrical and 20-60 μm long and 2-2.5 μm wide. The structure of the colony is circular (7mm in diameter after 14 days at 24 °C '' in vitro''), elevated and fuzzy in texture due to branched, filamentous growth. Large colonies can amalgamate into amorphous structures that optimizes the absorption of volatile compounds from the air, moving away from a more circular shapes and creating sheets of mycelium, especially under very humid conditions. '' In vitro'', ''Z. cellare'' is characterized as being morphologically similar to ''
Stenella araguata ''Stenella araguata'' is a species of anamorphic fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. Th ...
,'' which both reside in the order Capnodiales and family Mycosphaerellaceae.


Ecology

''Z. cellare'' was first recorded existing is cellars in 1696. Since that time it has not spread much, existing primarily in dark, humid, ethanol-rich cellars containing barrel aged wine, brandy or other spirits of central and southern Europe, Hungary, Poland, Great Britain and Nigeria. The presence of ''Z. cellare'' in any of these regions is completely dependent on human activity, however. For example, there are wineries in Italy that are devoid of ''Z. cellare'' but also maintain a level of cleanliness in their cellars, while other Italian wineries cherish its presence and often feed it left over wine, encouraging its proliferation. When ''Z. cellare'' is present, however, it is commonly found on brickwork and timber next to its food sources and can also, on the rare occasion, be found in nearby soil.


Physiology


Sexual physiology

The sexual state of ''Z. cellare'' is difficult to define, but there seems to be a common consensus in the literature that this fungus rarely, or never, presents itself as its
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 ...
. With that determined, the literature is ripe with contrary findings on what the proper state of the fungus is. Some argue that there is no optimal state, inherently defining it as either amorphous or in flux between anamorph and teleomorph, while others claim that it is all together
sterile Sterile or sterility may refer to: *Asepsis, a state of being free from biological contaminants * Sterile (archaeology), a sediment deposit which contains no evidence of human activity *Sterilization (microbiology), any process that eliminates or ...
, reproducing by
fragmentation Fragmentation or fragmented may refer to: Computers * Fragmentation (computing), a phenomenon of computer storage * File system fragmentation, the tendency of a file system to lay out the contents of files non-continuously * Fragmented distributi ...
. There is evidence, however, that ''Z. cellare'' produces
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s which could mean that it reproduces through asexual dispersal, but that is only a probable guess. What is known is that spore concentration is 2 times higher in cellars with ''Z. cellare'' than ones that do not. Further insights are awaited on this topic before the reproduction of this species is known, but this aspect of its life cycles makes ''Z. cellare'' unique among most other ''
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 ...
'' due to its difficulty to define by biology.


Mitochondrial genome

Apart from its reproduction, ''Z. cellare'' possesses an additional quality that, too, makes it distinct in its division. Of all
filamentous The word filament, which is descended from Latin ''filum'' meaning " thread", is used in English for a variety of thread-like structures, including: Astronomy * Galaxy filament, the largest known cosmic structures in the universe * Solar filament ...
ascomycete 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 defi ...
, ''Z. cellare'' contains the smallest known mitochondrial genome at 23 743
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
s, which is achieved by its
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
coding for proteins using smaller genes - lacking
intron An intron is any nucleotide sequence within a gene that is not expressed or operative in the final RNA product. The word ''intron'' is derived from the term ''intragenic region'', i.e. a region inside a gene."The notion of the cistron .e., gene. ...
s, non-essential genes and what is known as
noncoding Non-coding DNA (ncDNA) sequences are components of an organism's DNA that do not encode protein sequences. Some non-coding DNA is transcribed into functional non-coding RNA molecules (e.g. transfer RNA, microRNA, piRNA, ribosomal RNA, and regula ...
, or junk, DNA. Furthermore, there is an unusual feature in the sequence of this
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
consisting of a repeated 110
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
sequence that is inverted and separated by 1 000
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
s. As interesting as this is in the field of genetics, this is found to not be significant in any way.


Food consumption and energy production

Primarily, the food source of ''Z. cellare'' is ethanol from the process of barrel aging, but upon further investigation it was found that ''Z. cellare'' can survive, and even thrive, on much more. Chlebicki and Majewska (2010) discovered that this fungus can utilize any volatile, oxygen-containing organic compound including various other
alcohol Alcohol most commonly refers to: * Alcohol (chemistry), an organic compound in which a hydroxyl group is bound to a carbon atom * Alcohol (drug), an intoxicant found in alcoholic drinks Alcohol may also refer to: Chemicals * Ethanol, one of sev ...
s, esters,
acetic acid Acetic acid , systematically named ethanoic acid , is an acidic, colourless liquid and organic compound with the chemical formula (also written as , , or ). Vinegar is at least 4% acetic acid by volume, making acetic acid the main component ...
s, acetylaldehydes, as well as formaldehyde and thymol. Out of these compounds, it is found that alcohols and acids that are three to five carbons in length are preferred by this organism, but will gladly feed on any length of the former compounds if need be. As well, concentration is not a crucial factor by any means as the typical laboratory air concentration for volatile, organic compounds is enough for ''Z. cellare'' to grow. Given that alcohols are the fungus's main food source, it is likely that the organism produces energy via the citric acid cycle, where ethanol is converted to acetaldehyde, then to acetic acid, then to
acetyl-CoA Acetyl-CoA (acetyl coenzyme A) is a molecule that participates in many biochemical reactions in protein, carbohydrate and lipid metabolism. Its main function is to deliver the acetyl group to the citric acid cycle (Krebs cycle) to be oxidized for ...
via various enzymatic pathways which then joins the citric acid cycle to create energy for the organism from the oxidation of
acetyl-CoA Acetyl-CoA (acetyl coenzyme A) is a molecule that participates in many biochemical reactions in protein, carbohydrate and lipid metabolism. Its main function is to deliver the acetyl group to the citric acid cycle (Krebs cycle) to be oxidized for ...
. However, this is not proven.


Pathology

''Z. cellare'' has no recorded pathological effect on healthy individuals and has long been considered beneficial to human health by traditional, European winemakers who found a correspondence to the presence of this fungi and the elimination of musty odours. Schanderl (1950) saw it as proof that the volatile food sources of ''Z. cellare'' backed up this claim and too considered the presence of this fungus to be beneficial to human health. However, in unhealthy individuals such as English physician Sir John Floyer, the presence of ''Z. cellare'' had different consequences for him. In his essay ''A Treatise of the Asthma'' (1698), he mentions that being in the vicinity of a cellar, which at that time in Great Britain would likely be covered in ''Z. cellare'', will trigger asthma attacks. Though this is circumstantial evidence, it is an indication that ''Z. cellare'' might cause an immunological reaction in certain predisposed individuals that biology has yet to investigate.


References

{{Taxonbar, from=Q1734843 Fungi described in 1794 Taxa named by Christiaan Hendrik Persoon Mycosphaerellaceae Fungus species