''Coprinopsis cinerea'' is a species of

mushroom A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans. The standard for the name "mushroom" is t ...

in the family

Psathyrellaceae The Psathyrellaceae are a family of dark-spored agarics that generally have rather soft, fragile fruiting bodies, and are characterized by black, dark brown, rarely reddish, or even pastel-colored spore prints. About 50% of species produce fruit ...

. Commonly known as the gray shag, it is

edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...

, but must be used promptly after collecting. ''Coprinopsis cinerea'' is an important

model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...

for studying fungal sex and

mating type Mating types are the microorganism equivalent to sexes in multicellular lifeforms and are thought to be the ancestor to distinct Sex, sexes. They also occur in macro-organisms such as fungi. Definition Mating types are the microorganism equivalent ...

s, mushroom development, and the evolution of multicellularity of

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. These organisms are classified as a kingdom, separately from ...

. The

genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...

sequence was published in 2010. It is considered to be particularly suited organism to study

meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...

, due to its synchronous meiotic development and prolonged

prophase Prophase () is the first stage of cell division in both mitosis and meiosis. Beginning after interphase, DNA has already been replicated when the cell enters prophase. The main occurrences in prophase are the condensation of the chromatin retic ...

Research

Antibiotics

Researchers in 2014 discovered a protein produced by ''Coprinopsis cinerea'' with antibiotic properties. The protein, known as

copsin Copsin is a fungal defensin that acts as an antimicrobial polypeptide secreted from the inky cap mushroom, first reported at the end of 2014. The fungal defensin acts against gram positive bacteria. History In October 2014, a collaboration of t ...

, has similar effects to other non-protein organically derived antibiotics. To date, it has not been determined whether antibiotic medicine for humans and other animals can be developed from this protein.

Culturing

''Coprinopsis cinerea'' can be grown on complex (e.g. YMG, YMG/T) or minimal media (e.g. mKjalke medium), solid or liquid, with or without agitation, at 25 °C or optimally at 37 °C. It can be grown in dark or with 12-h light/12-h dark cycle.Efficient gene targeting in ΔCc.ku70 or ΔCc.lig4 mutants of the agaricomycete Coprinopsis cinerea.Lcc1 and Lcc5 are the main laccases secreted in liquid cultures of Coprinopsis cinerea strains

Strains

*C. cinereus strain PG78 (A6B42, trp1.1;1.6, pab1) is an AmutBmut monokaryon, self-compatible strain, with trp- and pab-auxotrophic markers (requires tryptophan and p-aminobenzoate).Restriction enzyme-mediated DNA integration in Coprinus cinereus

Genome

''Coprinopsis cinerea'' strain Okayama 7 (#130) was sequenced with 10x coverage in 2003. A third and most recent revision of the sequence of strain Okayama 7 (#130) was released in 2010. Its haploid genome is ca. 37.5 Mb.

Molecular cloning

''Coprinopsis cinerea'' can be transformed with exogenous DNA by transformation when the fungus is a protoplast. It was found that disrupting (knockout or RNAi silencing) ''

ku70 Ku70 is a protein that, in humans, is encoded by the ''XRCC6'' gene. Function Together, Ku70 and Ku80 make up the Ku heterodimer, which binds to DNA double-strand break ends and is required for the non-homologous end joining (NHEJ) pathway of ...

'' homologue can increase gene targeting via increased homologous recombination. Either protoplasts derived from oidia or vegetative mycelium can be used, however, gene targeting was found to be higher by 2% (based on phenotyping) when using vegetative mycelium. Otherwise, insertion of integrative vectors ectopically and with small homologous regions can be used, likely with low transformation efficiency. Earlier, REMI (restriction enzyme-mediated integration) could be used to insert exogenous DNA into the chromosome to produce mutant strains. This relies on inserting exogenous DNA and restriction enzymes into the protoplast cell, allowing for the enzymes to cut the chromosome at specific sites which match those sites used to produce linearized plasmid DNA with the gene of interest; subsequently, host enzymes ligate the cut sites and thus produce integrated heterologous, exogenous DNA. Although successful, undesirable mutations are likely. Chemical mutagenesis (also random) can also be done. Phenotype selection of the inability to fruit can indict that insertion led to disruption of vital genes. All in all, homologous recombination provides more specificity when creating a mutant strain. Depending on the mutant, auxotrophy markers (requires lost gene to be inserted) or prototrophy (when causing essential gene deletion) be used for selection.

Enzymes

''Coprinopsis cinerea'' is known to produce

laccase Laccases () are multicopper oxidases found in plants, fungi, and bacteria. Laccases oxidize a variety of phenolic substrates, performing one-electron oxidations, leading to crosslinking. For example, laccases play a role in the formation of lign ...

, a type of phenoloxidase. ''C. cinerea'' produces a variety of the same laccase, known as isoenzymes. Laccase activity can be measured by

zymogram Zymography is an Gel electrophoresis, electrophoretic technique for the detection of hydrolytic enzymes, based on the substrate repertoire of the enzyme. Three types of zymography are used; ''in gel'' zymography, ''in situ'' zymography and ''in vi ...

s (in which a substrate for the enzyme is present in a separating gel). Under stressed conditions, temperature and medium, laccase secretion was increased. Although copper is required co-factor for laccase, merely adding copper did not induce laccase secretion. It was recently found that a

TET Tet or TET may refer to: Vietnam *Tết or Tết Nguyên Đán, the Vietnamese new year, Lunar new year *Tet Offensive, a military campaign during the Vietnam War that began in 1968 **Tet 1969 Geography *Têt (river) in Roussillon, France *Tét, ...

(Ten-Eleven translocation dioxygenases) homologue, ''CcTET'', was identified in ''C. cinerea'', which may have important human (or mammalian) implications like cancer. DNA methylation is vital in humans and dysfunction is associated with cancer, thus, studying methylation reactions in non-mammalians may provide better insight into mammalian methylation reactions.

Reproduction

''Coprinopsis cinerea'' can sense blue light. It was identified that gene ''Cc.wc-2'' is involved in blue light photoreception. Etiolated stipes (elongation without cap maturation) is caused when grown without light.

Meiosis

''Coprinopsis cinerea'' is an ideal model for studying meiosis because meiosis progresses synchronously in about 10 million cells within each mushroom cap. Meiosis is a specialized cell division process, occurring in diploid cells, in which a single round of DNA replication occurs, and is followed by two divisions to produce four haploid daughter nuclei. During meiosis homologous chromosomes pair with each other and undergo a DNA repair process in which DNA damage is removed and genetic information is recombined. Burns et al. studied the expression of genes involved in the 15-hour meiotic process encompassing time points prior to the haploid nuclear fusion that forms the diploid zygote to the final formation of the four haploid products. They compared expression of particular genes in ''C. cinerea'' to the expression of the comparable genes (orthologs) in two other species (''

Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...

'' and ''

Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically meas ...

'') from which ''C. cinerea'' had diverged in evolution 500 to 900 million years ago. They found that the expression of individual genes turned on or off at the same stage in ''C. cinerea'' as in the other two species. They also found that genes considered to be specifically involved in the meiotic process were more conserved in their expression pattern than non-meiotic genes. These findings indicate ancient conservation of the meiotic process.

Human disease

''Coprinopsis cinerea'' is harmless to human and animal health under normal conditions. However, the organism can cause opportunistic infections (

mycoses Fungal infection, also known as mycosis, is 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 ti ...

) in

immunocompromised Immunodeficiency, also known as immunocompromisation, is a state in which the immune system's ability to fight infectious diseases and cancer is compromised or entirely absent. Most cases are acquired ("secondary") due to extrinsic factors that a ...

patients, such as those who have undergone haematopoietic stem cell transplantation or are otherwise undergoing immunosuppression treatment. Most reported cases have been respiratory infections, but cases involving the heart, skin, brain or gut have been reported, and the infections may rapidly become systemic. Whilst exceptionally rare, ''Coprinopsis cinerea'' infection is difficult to treat and often fatal in this vulnerable patient group. The infection is caused by the mould-like asexual (non mushroom-forming)

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 ...

of ''Coprinopsis cinerea'' which used to be known as ''Hormographiella aspergillata'', and may be described under this name in the clinical literature.Correa-Martinez C, Brentrup A, Hess K, Becker K, Groll AH, Schaumburg F. First description of a local Coprinopsis cinerea skin and soft tissue infection. New microbes and new infections. 2018 Jan 1;21:102-4.

References

{{Taxonbar, from=Q5168926 cinerea Edible fungi Fungi of Asia Fungi of Europe Fungi of North America Fungi described in 1774 Ammonia fungi Taxa named by Jacob Christian Schäffer