Coprinellus micaceus
Encyclopedia
Coprinellus micaceus is a common species of fungus
in the family Psathyrellaceae
with a cosmopolitan distribution
. The fruit bodies
of the saprobe typically grow in clusters on or near rotting hardwood
tree stumps or underground tree roots. Depending on their stage of development, the tawny
-brown mushroom caps
may range in shape from oval to bell-shaped to convex, and reach diameters up to 3 cm (1.2 in). The caps, marked with fine radial grooves that extend nearly to the center, rest atop whitish stems
up to 10 cm (3.9 in) long. In young specimens, the entire cap surface is coated with a fine layer of reflective mica
-like cells that provide the inspiration for both the mushroom's species name and the common name
s mica cap, shiny cap, and glistening inky cap. Although small and with thin flesh
, the mushrooms are usually bountiful, as they typically grow in dense clusters. A few hours after collection, the gills will begin to slowly dissolve into a black, inky, spore
-laden liquid—an enzymatic
process called autodigestion or deliquescence. The fruit bodies are edible
before the gills blacken and dissolve, and cooking will stop the autodigestion process.
The microscopic characteristics and cytogenetics
of C. micaceus are well-known, and it has been used frequently as a model organism
to study cell division
and meiosis
in Basidiomycetes. Chemical analysis of the fruit bodies has revealed the presence of antibacterial and enzyme
-inhibiting compounds. Formerly known as Coprinus micaceus, the species was transferred to Coprinellus
in 2001 as phylogenetic analyses provided the impetus for a reorganization of the many species formerly grouped together in the genus Coprinus
. Based on external appearance, C. micaceus is virtually indistinguishable from C. truncorum
, and it has been suggested that many reported collections of the former may be of the latter.
by the 16th-century botanist Carolus Clusius in what is arguably the first published monograph
on fungi, the 1601 Rariorum plantarum historia. Fungorum in Pannoniis observatorum brevis historia (History of rare plants. Brief history of fungi observed in Pannonia [Hungary]). Clusius erroneously believed the species to be poisonous, and classified it as a genus of Fungi perniciales (harmful fungi). The species was first described scientifically by French botanist Jean Baptiste François Pierre Bulliard
in 1786 as Agaricus micaceus in his work Herbier de la France. In 1801, Christian Hendrik Persoon
grouped together all of the gilled fungi that auto-digested (deliquesced) during spore discharge into the section Coprinus of the genus Agaricus. Elias Magnus Fries
later raised Persoon's section Coprinus to genus rank in his Epicrisis Systematis Mycologici, and the species became known as Coprinus micaceus. It was the type species
of subsection Exannulati in section Micacei of the genus Coprinus, a grouping of related taxa
with veils
made of sphaerocysts (round swollen cells usually formed in clusters) exclusively or with thin-filamentous connective hypha
e intermixed. Molecular studies published in the 1990s demonstrated that many of the coprinoid (Coprinus-like) mushrooms were in fact unrelated to each other. This culminated in a 2001 revision of the genus Coprinus
, which was split into four genera; C. micaeus was transferred to Coprinellus
.
Due partly to their ready availability and the ease with which they may be grown in the laboratory, C. micaceus and other coprinoid mushrooms were common subjects in cytological
studies of the 19th and 20th centuries. The German botanist Johann Heinrich Friedrich Link
reported his observations of the structure of the hymenium
(the fertile spore
-bearing surface) in 1809, but misinterpreted what he had seen. Link thought that microscopic structures known today as basidia were thecae, comparable in form to the asci
of the Ascomycetes, and that each theca contained four series of spores. His inaccurate drawings of the hymenium of C. micaceus were copied in subsequent mycological
publications by other authors, and it was not until microscopy had advanced that mycologists were able to determine the true nature of the basidia, when nearly three decades later in 1837 Joseph-Henri Léveillé
and August Corda independently published correct descriptions of the structure of the hymenium. In 1924, A. H. Reginald Buller published a comprehensive description and analysis of the processes of spore production and release in the third volume of his Researches on Fungi.
The specific epithet micaceus is derived from the Latin
word mica, for "crumb, grain of salt" and the suffix -aceus, "like, similar"; the modern application of "mica" to a very different substance
comes from the influence of micare, "glitter". The mushroom is commonly
known as the "shiny cap", the "mica cap" or the "glistening inky cap", all in reference to the mealy particles found on the cap that glisten like mica
.
is initially 1 – in diameter, oval to cylindrical, but expands to become campanulate (bell-shaped), sometimes with an umbo
(a central nipple-like protrusion); finally it flattens somewhat, becoming convex. When expanded, the cap diameter reaches 0.8 – with the margin torn into rays and turned upwards slightly. The color is yellow-brown or tan often with a darker center, then pale yellow or buff from the margin inwards. The cap margin is prominently grooved almost all the way to the center; the grooves mark the positions of the longer gills on the underside of the cap. When young, the cap surface is covered with white or whitish shiny particles, remnants of the universal veil
that covers immature specimens. The particles are loosely attached and easily washed away, so that older specimens are often smooth. Coprinellus micaceus is hygrophanous
, meaning it assumes different colors depending on its state of hydration.
The gills are crowded together closely, and have an adnexed (narrow) attachment to the stem
. Initially white, they change color to dark brown then eventually black as the spores mature. Expansion of the cap causes the gills to split open down their median planes, tearing the cap margin into rays. The process of spore discharge and autodigestion begin at the bottom of the gills before the upper parts of the gills have become completely blackened. The brittle stem is hollow, and measures 4 – long by 0.2 – thick and is roughly the same diameter throughout the length of the stem. It is generally white, but may discolor to pale dirty cream from the base up. The stem surface is at first velvety with a very fine whitish powder, but this eventually wears off, leaving it more or less smooth. Stems may have a rudimentary ring at the base, another universal veil remnant. The spore print is dark brown or black. The flesh
is thin, fragile, white in the stem, and brownish in the cap. Its odor and taste are not distinctive. Individual fruit bodies take an average of five to seven days to fully mature.
. Generally, they are lentiform (shaped like a biconvex lens), but viewed from the side they appear more almond-shaped or spindle-shaped, while in front view they appear oval or mitriform (roughly the shape of a miter—a peaked cap). Spores have a germ pore
, a flattened area in the center of the spore surface through which a germ tube
may emerge. The spore-bearing cells (the basidia) are four-spored, club-shaped, and measure 10–15 by 4–7 µm. Studies have shown that the basidia develop in four discrete generations. The first generation basidia are the most protuberant, and extend out the greatest distance from the surface of the hymenium. Subsequent generations of basidia have shorter and less protuberant bodies. When a living gill is viewed with a microscope, the four sets of basidia can be seen distinctly. Arthur Buller
coined the term inaequihymeniiferous to describe this mode of hymenial development. The purpose of the staggered basidia sizes is to facilitate the release of spores from the hymenium. There are four zones of spore discharge that correspond to the four sets of basidia, and basidia that have released all of their spores quickly begin to autodigest. The staggered setup minimizes the chance of spores colliding with neighboring basidia during release.
Cystidia that are located along the edge of the cap (called cheilocystidia) are spherical, and 30–120 by 20–74 µm. The facial cystidia (called pleurocystidia) are club-shaped or elongated ellipses, up to 130–155 µm in length. The pleurocystidia protrude from the face of the gill and act as guards, preventing adjacent gills from touching each other, and also ensuring that the basidia and spores have sufficient room for development. C. micaceus may also have scattered caulocystidia (cystidia on the stem) that are 60–100 by 5–10 µm, but their presence is variable and cannot reliably be used for identification. Both De Bary and Buller, in their investigations into the structure of the cystidia, concluded that there is a central mass of cytoplasm
formed where numerous thin plates of cytoplasm meet at the center of the cell. De Bary believed that the plates were filamentous branching processes, but Buller thought that they were formed in a process similar to the walls of foam bubbles, and that the central mass was able to slowly change form and position by altering the relative volumes of the vacuole
s enclosed by the numerous thin cytoplasmic walls. In older cells, the cytoplasm may be limited to the periphery of the cell, with one huge vacuole occupying the cell center.
The globular cells that make up the mica-resembling scales on the cap are colorless, smooth-walled, and range in size from about 25–65 µm, although most are between 40–50 µm. Buller explained the "glitter" of these cells as follows: "The sparkling of the meal-cells, as well as of the cystidia on the edges and faces of the gills, is simply due to light which strikes them from without and is refracted and reflected to the eye in the same manner as from the minute drops of water one so often sees at the tips of grass leaves on English lawns early in the morning after a dewy night."
In 1914, Michael Levine was the first to report successfully cultivating Coprinopsis micaceus from spores in the laboratory. In his experiments, fruit bodies appeared roughly 40 to 60 days after initially inoculating the growth media (agar
supplemented with soil, horse dung, or cornmeal
) with spores. Like other coprinoid species, C. micaceus undergoes synchronous meiosis
. The chromosome
s are readily discernible with light microscopy, and all of the meiotic stages are well-defined. These features have made the species a useful tool in laboratory investigations of Basidiomycete cytogenetics
. The chromosome number of C. micaceus is n=12.
s that cause autodigestion or deliquescence—a process that can begin as soon as one hour after collection. It is considered ideal for omelettes, and as a flavor for sauces, although it is "a very delicate species easily spoiled by overcooking". The fungus also appeals to fruit flies of the genus Drosophila
, who frequently use the fruit bodies as hosts for larva
e production.
A study of the mineral
contents of various edible mushrooms found that C. micaceus contained the highest concentration of potassium
in the 34 species tested, close to half a gram of potassium per kilogram of mushroom. Because the species can bioaccumulate
detrimental heavy metals like lead
and cadmium
, it has been advised to restrict consumption of specimens collected from roadsides or other collection sites that may be exposed to or contain pollutants.
is nearly identical but lacks the yellowish cap granules and only has two spores per basidium. The scaly inky cap (Coprinus variegatus = Coprinus quadrifidus) has a grayish-brown cap with dull white to brownish scales; its odor is disagreeable. The trooping crumble cap (Coprinellus disseminatus
, edible) has smaller, yellow-brown to grey-brown caps and white gills that turn black but do not dissolve away; it always grows in large clusters on rotting wood (sometimes buried wood). Coprinus atramentarius is a larger, gray species that grows in dense clusters on stumps or on the ground from buried wood, lacks glistening particles on the cap, and the cap and gills dissolve at maturity. Coprinellus radians
develops singly or in clumps from a tufted mat of coarse yellow-orange mycelium
on the wood. C. truncorum is also covered with glistening granules and is said to be almost indistinguishable from C. micaceus in the field; microscopy is needed to tell the difference, as C. truncorum has ellipsoid spores with a rounded germ pore, compared to the shield-shaped (mitriform) spores with truncated germ pores of C. micaceus. One study suggests that compared to C. truncorum, C. micaceus is browner in the center of the cap (rather than grayish) and has a greater tendency to grow in clusters; more molecular
evidence is required to determine if the two taxa are genetically identical.
, and grows in and around stumps or logs of broad-leaved trees or attached to buried wood. It prefers feeding on bark
, particularly the secondary phloem, rather than the wood. In the scheme of the succession
of fungal species involved in the decomposition of wood, C. micaceus is a late stage colonizer, and prefers to feed on wood that has already decomposed sufficiently to have reached "a friable softened consistency". A 2010 study suggests that the fungus can also live as an endophyte
, inhabiting the woody tissue of healthy trees without causing disease symptoms. The fungus is also associated with disturbed or developed ground, such as the sides of roads and paths, gardens, building sites and the edges of parking lots; it has also been noted for growing indoors on rotting wood in humid environments. In one instance it was discovered about 120 m (400 ft) underground in an abandoned coal mine, growing on wooden gangways and props used to support the roof.
Fruit bodies are commonly found growing in dense clusters, but can also be found growing singly or in small clumps, especially in forested areas. In North America, C. micaceus is one of the first edible mushrooms to appear in the spring, and fruits from May to September. In Europe, it fruits from May to December. Although it can grow at any time of the year, it is more prevalent during the spring and fall, coinciding with the higher humidity resulting from spring and autumn rains. A study of air quality conducted in the city of Santiago de Compostela
in the Iberian Peninsula
, concluded that most "Coprinus" spores present in the atmosphere belonged to C. micaceus, and that the number of spores went up with increased humidity
and rainfall, but decreased with greater temperatures. The species is known for reappearing with successive fruitings at the same location. In one case, a total of 38 lb (17.2 kg) of fresh mushrooms were collected from one elm
stump in 10 successive crops over a spring and summer.
Coprinellus micaceus has a cosmopolitan distribution
, and has been collected in northern Africa, South Africa, Europe (including Turkey
), North America (as far north as Alaska
), the Hawaiian islands, South America, India, Australia, New Zealand, and Japan. Phylogenetic analysis of rDNA
sequences from specimens collected in southeastern Asia and Hawaii show that the Hawaiian species form a distinct clade
with little genetic diversity
compared to Asian populations; this suggests that the Hawaiian populations have been introduced relatively recently and have not had much time to develop genetic variation. One study suggests that in South Africa, where C. micaceus is rare, it has been frequently confused with the similar-appearing C. truncorum
, a more common species in that region. A similar inference has been raised about North American species.
chemistry of Coprinellus micaceus has revealed the presence of several chemical compounds unique to the species. Micaceol is a sterol
with "modest" antibacterial activity against the pathogens Corynebacterium xerosis and Staphylococcus aureus
. The compound (Z,Z)-4-oxo-2,5-heptadienedioic acid has inhibitory activity against glutathione S-transferase
, an enzyme
that has been implicated in the resistance of cancer cells against chemotherapeutic
agents, especially alkylating
drugs. A 2003 study did not find any antibacterial activity in this species. A 1962 publication reported the presence of the biologically active indole
compound tryptamine
in C. micaceus, although the concentration was not determined. The fruit bodies additionally produce a variety of pigment
compounds known as melanin
s—complex chemical polymer
s that contribute to the formation of soil humus after the fruit bodies have disintegrated. C. micaceus has been found to be devoid of the toxin coprine, the Antabuse-mimicking chemical found in Coprinus comatus
that causes illness when consumed simultaneously with alcohol.
Fungus
A fungus is a member of a large 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, Fungi, which is separate from plants, animals, and bacteria...
in the family Psathyrellaceae
Psathyrellaceae
The Psathyrellaceae are a family of dark-spored agarics that generally have rather soft, fragile fruiting bodies, and are characterized by black or dark brown, rarely reddish, or even pastel colored spore prints. About 50% of the species produce fruiting bodies that dissolve into ink-like ooze...
with a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...
. The fruit bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
of the saprobe typically grow in clusters on or near rotting hardwood
Hardwood
Hardwood is wood from angiosperm trees . It may also be used for those trees themselves: these are usually broad-leaved; in temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen.Hardwood contrasts with softwood...
tree stumps or underground tree roots. Depending on their stage of development, the tawny
Tawny (color)
Tawny is a yellowish brown color. The word means "tan-colored," from Anglo-Norman tauné "associated with the brownish-yellow of tanned leather," from Old French tané "to tan hides," from Medieval Latin tannare, from tannum "crushed oak bark," used in tanning leather, probably from a Celtic source...
-brown mushroom caps
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
may range in shape from oval to bell-shaped to convex, and reach diameters up to 3 cm (1.2 in). The caps, marked with fine radial grooves that extend nearly to the center, rest atop whitish stems
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
up to 10 cm (3.9 in) long. In young specimens, the entire cap surface is coated with a fine layer of reflective mica
Mica
The mica group of sheet silicate minerals includes several closely related materials having highly perfect basal cleavage. All are monoclinic, with a tendency towards pseudohexagonal crystals, and are similar in chemical composition...
-like cells that provide the inspiration for both the mushroom's species name and the common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
s mica cap, shiny cap, and glistening inky cap. Although small and with thin flesh
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....
, the mushrooms are usually bountiful, as they typically grow in dense clusters. A few hours after collection, the gills will begin to slowly dissolve into a black, inky, spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
-laden liquid—an enzymatic
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
process called autodigestion or deliquescence. The fruit bodies are edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...
before the gills blacken and dissolve, and cooking will stop the autodigestion process.
The microscopic characteristics and cytogenetics
Cytogenetics
Cytogenetics is a branch of genetics that is concerned with the study of the structure and function of the cell, especially the chromosomes. It includes routine analysis of G-Banded chromosomes, other cytogenetic banding techniques, as well as molecular cytogenetics such as fluorescent in situ...
of C. micaceus are well-known, and it has been used frequently as a model organism
Model organism
A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the organism model will provide insight into the workings of other organisms. Model organisms are in vivo models and are widely used to...
to study cell division
Cell division
Cell division is the process by which a parent cell divides into two or more daughter cells . Cell division is usually a small segment of a larger cell cycle. This type of cell division in eukaryotes is known as mitosis, and leaves the daughter cell capable of dividing again. The corresponding sort...
and meiosis
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....
in Basidiomycetes. Chemical analysis of the fruit bodies has revealed the presence of antibacterial and enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
-inhibiting compounds. Formerly known as Coprinus micaceus, the species was transferred to Coprinellus
Coprinellus
Coprinellus is a genus of mushrooms in the family Psathyrellaceae. The genus was first described by Petter Karsten in 1879.-External links:* at Index Fungorum...
in 2001 as phylogenetic analyses provided the impetus for a reorganization of the many species formerly grouped together in the genus Coprinus
Coprinus
Coprinus is a small genus of mushrooms consisting of Coprinus comatus and several of its close relatives. Until 2001, Coprinus was a large genus consisting of all agaric species in which the lamellae autodigested to release their spores...
. Based on external appearance, C. micaceus is virtually indistinguishable from C. truncorum
Coprinellus truncorum
Coprinellus truncorum is a species of mushroom in the Psathyrellaceae family. Part of the cluster of mushroom morphologically related to Coprinellus micaceus, this species can be distinguished from C. micaceus by a smooth, rather than pruinose stem, and by having more elliptical spores...
, and it has been suggested that many reported collections of the former may be of the latter.
History and taxonomy
Coprinellus micaeus was illustrated in a woodcutWoodcut
Woodcut—occasionally known as xylography—is a relief printing artistic technique in printmaking in which an image is carved into the surface of a block of wood, with the printing parts remaining level with the surface while the non-printing parts are removed, typically with gouges...
by the 16th-century botanist Carolus Clusius in what is arguably the first published monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...
on fungi, the 1601 Rariorum plantarum historia. Fungorum in Pannoniis observatorum brevis historia (History of rare plants. Brief history of fungi observed in Pannonia [Hungary]). Clusius erroneously believed the species to be poisonous, and classified it as a genus of Fungi perniciales (harmful fungi). The species was first described scientifically by French botanist Jean Baptiste François Pierre Bulliard
Jean Baptiste François Pierre Bulliard
Jean Baptiste François Pierre Bulliard was a French physician and botanist....
in 1786 as Agaricus micaceus in his work Herbier de la France. In 1801, Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...
grouped together all of the gilled fungi that auto-digested (deliquesced) during spore discharge into the section Coprinus of the genus Agaricus. Elias Magnus Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...
later raised Persoon's section Coprinus to genus rank in his Epicrisis Systematis Mycologici, and the species became known as Coprinus micaceus. It was the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
of subsection Exannulati in section Micacei of the genus Coprinus, a grouping of related taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
with veils
Partial veil
thumb|150px|right|Developmental stages of [[Agaricus campestris]] showing the role and evolution of a partial veilPartial veil is a mycological term used to describe a temporary structure of tissue found on the fruiting bodies of some basidiomycete fungi, typically agarics...
made of sphaerocysts (round swollen cells usually formed in clusters) exclusively or with thin-filamentous connective hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e intermixed. Molecular studies published in the 1990s demonstrated that many of the coprinoid (Coprinus-like) mushrooms were in fact unrelated to each other. This culminated in a 2001 revision of the genus Coprinus
Coprinus
Coprinus is a small genus of mushrooms consisting of Coprinus comatus and several of its close relatives. Until 2001, Coprinus was a large genus consisting of all agaric species in which the lamellae autodigested to release their spores...
, which was split into four genera; C. micaeus was transferred to Coprinellus
Coprinellus
Coprinellus is a genus of mushrooms in the family Psathyrellaceae. The genus was first described by Petter Karsten in 1879.-External links:* at Index Fungorum...
.
Due partly to their ready availability and the ease with which they may be grown in the laboratory, C. micaceus and other coprinoid mushrooms were common subjects in cytological
Cell biology
Cell biology is a scientific discipline that studies cells – their physiological properties, their structure, the organelles they contain, interactions with their environment, their life cycle, division and death. This is done both on a microscopic and molecular level...
studies of the 19th and 20th centuries. The German botanist Johann Heinrich Friedrich Link
Johann Heinrich Friedrich Link
Johann Heinrich Friedrich Link was a German naturalist and botanist.Link was born at Hildesheim as a son of the minister August Heinrich Link , who taught him the love for nature through collection of 'natural objects'...
reported his observations of the structure of the hymenium
Hymenium
The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some cells develop into sterile cells called cystidia or...
(the fertile spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
-bearing surface) in 1809, but misinterpreted what he had seen. Link thought that microscopic structures known today as basidia were thecae, comparable in form to the asci
Ascus
An ascus is the sexual spore-bearing cell produced in ascomycete fungi. On average, asci normally contain eight ascospores, produced by a meiotic cell division followed, in most species, by a mitotic cell division. However, asci in some genera or species can number one , two, four, or multiples...
of the Ascomycetes, and that each theca contained four series of spores. His inaccurate drawings of the hymenium of C. micaceus were copied in subsequent mycological
Mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans as a source for tinder, medicinals , food and entheogens, as well as their dangers, such as poisoning or...
publications by other authors, and it was not until microscopy had advanced that mycologists were able to determine the true nature of the basidia, when nearly three decades later in 1837 Joseph-Henri Léveillé
Joseph-Henri Léveillé
Joseph-Henri Léveillé was a French physician and mycologist who was a native of Crux-la-Ville, in the department of Nièvre....
and August Corda independently published correct descriptions of the structure of the hymenium. In 1924, A. H. Reginald Buller published a comprehensive description and analysis of the processes of spore production and release in the third volume of his Researches on Fungi.
The specific epithet micaceus is derived from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
word mica, for "crumb, grain of salt" and the suffix -aceus, "like, similar"; the modern application of "mica" to a very different substance
Mica
The mica group of sheet silicate minerals includes several closely related materials having highly perfect basal cleavage. All are monoclinic, with a tendency towards pseudohexagonal crystals, and are similar in chemical composition...
comes from the influence of micare, "glitter". The mushroom is commonly
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
known as the "shiny cap", the "mica cap" or the "glistening inky cap", all in reference to the mealy particles found on the cap that glisten like mica
Mica
The mica group of sheet silicate minerals includes several closely related materials having highly perfect basal cleavage. All are monoclinic, with a tendency towards pseudohexagonal crystals, and are similar in chemical composition...
.
Description
The capPileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
is initially 1 – in diameter, oval to cylindrical, but expands to become campanulate (bell-shaped), sometimes with an umbo
Umbo (mycology)
thumb|right|[[Cantharellula umbonata]] has an umbo.thumb|right|The cap of [[Psilocybe makarorae]] is acutely papillate.An umbo is a raised area in the center of a mushroom cap. Caps that possess this feature are called umbonate. Umbos that are sharply pointed are called acute, while those that are...
(a central nipple-like protrusion); finally it flattens somewhat, becoming convex. When expanded, the cap diameter reaches 0.8 – with the margin torn into rays and turned upwards slightly. The color is yellow-brown or tan often with a darker center, then pale yellow or buff from the margin inwards. The cap margin is prominently grooved almost all the way to the center; the grooves mark the positions of the longer gills on the underside of the cap. When young, the cap surface is covered with white or whitish shiny particles, remnants of the universal veil
Universal veil
In mycology, a universal veil is a temporary membranous tissue that fully envelops immature fruiting bodies of certain gilled mushrooms. The developing Caesar's mushroom , for example, which may resemble a small white sphere at this point, is protected by this structure...
that covers immature specimens. The particles are loosely attached and easily washed away, so that older specimens are often smooth. Coprinellus micaceus is hygrophanous
Hygrophanous
The adjective hygrophanous refers to the color change of mushroom tissue as it loses or absorbs water, which causes the pileipellis to become more transparent when wet and opaque when dry....
, meaning it assumes different colors depending on its state of hydration.
The gills are crowded together closely, and have an adnexed (narrow) attachment to the stem
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
. Initially white, they change color to dark brown then eventually black as the spores mature. Expansion of the cap causes the gills to split open down their median planes, tearing the cap margin into rays. The process of spore discharge and autodigestion begin at the bottom of the gills before the upper parts of the gills have become completely blackened. The brittle stem is hollow, and measures 4 – long by 0.2 – thick and is roughly the same diameter throughout the length of the stem. It is generally white, but may discolor to pale dirty cream from the base up. The stem surface is at first velvety with a very fine whitish powder, but this eventually wears off, leaving it more or less smooth. Stems may have a rudimentary ring at the base, another universal veil remnant. The spore print is dark brown or black. The flesh
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....
is thin, fragile, white in the stem, and brownish in the cap. Its odor and taste are not distinctive. Individual fruit bodies take an average of five to seven days to fully mature.
Microscopic characteristics
The spores of C. micaceus are reddish-brown, with dimensions of 7–10 by 4.5–6 µmMicrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
. Generally, they are lentiform (shaped like a biconvex lens), but viewed from the side they appear more almond-shaped or spindle-shaped, while in front view they appear oval or mitriform (roughly the shape of a miter—a peaked cap). Spores have a germ pore
Germ pore
A germ pore is a small pore in the outer wall of a fungal spore through which the germ tube exits upon germination. It can be apical or eccentric in its location, and, on light microscopy, may be visualized as a lighter coloured area on the cell wall....
, a flattened area in the center of the spore surface through which a germ tube
Germ tube
A germ tube is an outgrowth produced by spores of spore-releasing fungi during germination.The germ tube differentiates, grows, and develops by mitosis to create somatic hyphae....
may emerge. The spore-bearing cells (the basidia) are four-spored, club-shaped, and measure 10–15 by 4–7 µm. Studies have shown that the basidia develop in four discrete generations. The first generation basidia are the most protuberant, and extend out the greatest distance from the surface of the hymenium. Subsequent generations of basidia have shorter and less protuberant bodies. When a living gill is viewed with a microscope, the four sets of basidia can be seen distinctly. Arthur Buller
Arthur Henry Reginald Buller
Arthur Henry Reginald Buller was a British-Canadian mycologist. He is mainly known as a researcher of fungi and wheat rust.- Academic career :...
coined the term inaequihymeniiferous to describe this mode of hymenial development. The purpose of the staggered basidia sizes is to facilitate the release of spores from the hymenium. There are four zones of spore discharge that correspond to the four sets of basidia, and basidia that have released all of their spores quickly begin to autodigest. The staggered setup minimizes the chance of spores colliding with neighboring basidia during release.
Cystidia that are located along the edge of the cap (called cheilocystidia) are spherical, and 30–120 by 20–74 µm. The facial cystidia (called pleurocystidia) are club-shaped or elongated ellipses, up to 130–155 µm in length. The pleurocystidia protrude from the face of the gill and act as guards, preventing adjacent gills from touching each other, and also ensuring that the basidia and spores have sufficient room for development. C. micaceus may also have scattered caulocystidia (cystidia on the stem) that are 60–100 by 5–10 µm, but their presence is variable and cannot reliably be used for identification. Both De Bary and Buller, in their investigations into the structure of the cystidia, concluded that there is a central mass of cytoplasm
Cytoplasm
The cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the cytoplasm...
formed where numerous thin plates of cytoplasm meet at the center of the cell. De Bary believed that the plates were filamentous branching processes, but Buller thought that they were formed in a process similar to the walls of foam bubbles, and that the central mass was able to slowly change form and position by altering the relative volumes of the vacuole
Vacuole
A vacuole is a membrane-bound organelle which is present in all plant and fungal cells and some protist, animal and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic molecules including enzymes in solution, though in certain...
s enclosed by the numerous thin cytoplasmic walls. In older cells, the cytoplasm may be limited to the periphery of the cell, with one huge vacuole occupying the cell center.
The globular cells that make up the mica-resembling scales on the cap are colorless, smooth-walled, and range in size from about 25–65 µm, although most are between 40–50 µm. Buller explained the "glitter" of these cells as follows: "The sparkling of the meal-cells, as well as of the cystidia on the edges and faces of the gills, is simply due to light which strikes them from without and is refracted and reflected to the eye in the same manner as from the minute drops of water one so often sees at the tips of grass leaves on English lawns early in the morning after a dewy night."
In 1914, Michael Levine was the first to report successfully cultivating Coprinopsis micaceus from spores in the laboratory. In his experiments, fruit bodies appeared roughly 40 to 60 days after initially inoculating the growth media (agar
Agar
Agar or agar-agar is a gelatinous substance derived from a polysaccharide that accumulates in the cell walls of agarophyte red algae. Throughout history into modern times, agar has been chiefly used as an ingredient in desserts throughout Asia and also as a solid substrate to contain culture medium...
supplemented with soil, horse dung, or cornmeal
Cornmeal
Cornmeal is flour ground from dried maize or American corn. It is a common staple food, and is ground to fine, medium, and coarse consistencies. In the United States, the finely ground cornmeal is also referred to as cornflour. However, the word cornflour denotes cornstarch in recipes from the...
) with spores. Like other coprinoid species, C. micaceus undergoes synchronous meiosis
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....
. The chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...
s are readily discernible with light microscopy, and all of the meiotic stages are well-defined. These features have made the species a useful tool in laboratory investigations of Basidiomycete cytogenetics
Cytogenetics
Cytogenetics is a branch of genetics that is concerned with the study of the structure and function of the cell, especially the chromosomes. It includes routine analysis of G-Banded chromosomes, other cytogenetic banding techniques, as well as molecular cytogenetics such as fluorescent in situ...
. The chromosome number of C. micaceus is n=12.
Edibility
Coprinellus micaceus is an edible species, and cooking inactivates the enzymeEnzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
s that cause autodigestion or deliquescence—a process that can begin as soon as one hour after collection. It is considered ideal for omelettes, and as a flavor for sauces, although it is "a very delicate species easily spoiled by overcooking". The fungus also appeals to fruit flies of the genus Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
, who frequently use the fruit bodies as hosts for larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e production.
A study of the mineral
Dietary mineral
Dietary minerals are the chemical elements required by living organisms, other than the four elements carbon, hydrogen, nitrogen, and oxygen present in common organic molecules. Examples of mineral elements include calcium, magnesium, potassium, sodium, zinc, and iodine...
contents of various edible mushrooms found that C. micaceus contained the highest concentration of potassium
Potassium
Potassium is the chemical element with the symbol K and atomic number 19. Elemental potassium is a soft silvery-white alkali metal that oxidizes rapidly in air and is very reactive with water, generating sufficient heat to ignite the hydrogen emitted in the reaction.Potassium and sodium are...
in the 34 species tested, close to half a gram of potassium per kilogram of mushroom. Because the species can bioaccumulate
Bioaccumulation
Bioaccumulation refers to the accumulation of substances, such as pesticides, or other organic chemicals in an organism. Bioaccumulation occurs when an organism absorbs a toxic substance at a rate greater than that at which the substance is lost...
detrimental heavy metals like lead
Lead
Lead is a main-group element in the carbon group with the symbol Pb and atomic number 82. Lead is a soft, malleable poor metal. It is also counted as one of the heavy metals. Metallic lead has a bluish-white color after being freshly cut, but it soon tarnishes to a dull grayish color when exposed...
and cadmium
Cadmium
Cadmium is a chemical element with the symbol Cd and atomic number 48. This soft, bluish-white metal is chemically similar to the two other stable metals in group 12, zinc and mercury. Similar to zinc, it prefers oxidation state +2 in most of its compounds and similar to mercury it shows a low...
, it has been advised to restrict consumption of specimens collected from roadsides or other collection sites that may be exposed to or contain pollutants.
Similar species
The edible Coprinellus bisporusCoprinellus bisporus
Coprinellus bisporus is a species of mushroom in the Psathyrellaceae family. It was first described as Coprinus bisporus by mycologist Jakob Emanuel Lange in 1915, and later transferred to the genus Coprinellus in 2001 on the basis of phylogenetic analysis.-External Links:...
is nearly identical but lacks the yellowish cap granules and only has two spores per basidium. The scaly inky cap (Coprinus variegatus = Coprinus quadrifidus) has a grayish-brown cap with dull white to brownish scales; its odor is disagreeable. The trooping crumble cap (Coprinellus disseminatus
Coprinellus disseminatus
Coprinellus disseminatus is a species of mushroom in the Psathyrellaceae family. Unlike most other Coprinoid mushrooms, C. disseminatus does not dissolve into black ink in maturity. The species was given its current name in 1939 by Jakob Emanuel Lange...
, edible) has smaller, yellow-brown to grey-brown caps and white gills that turn black but do not dissolve away; it always grows in large clusters on rotting wood (sometimes buried wood). Coprinus atramentarius is a larger, gray species that grows in dense clusters on stumps or on the ground from buried wood, lacks glistening particles on the cap, and the cap and gills dissolve at maturity. Coprinellus radians
Coprinellus radians
Coprinellus radians is a species of mushroom in the Psathyrellaceae family. First described as Agaricus radians by the mycologist John Baptiste Henri Joseph Desmazières in 1828, it was later transferred to the genus Coprinellus in 2001.-See also:...
develops singly or in clumps from a tufted mat of coarse yellow-orange mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...
on the wood. C. truncorum is also covered with glistening granules and is said to be almost indistinguishable from C. micaceus in the field; microscopy is needed to tell the difference, as C. truncorum has ellipsoid spores with a rounded germ pore, compared to the shield-shaped (mitriform) spores with truncated germ pores of C. micaceus. One study suggests that compared to C. truncorum, C. micaceus is browner in the center of the cap (rather than grayish) and has a greater tendency to grow in clusters; more molecular
Molecular genetics
Molecular genetics is the field of biology and genetics that studies the structure and function of genes at a molecular level. The field studies how the genes are transferred from generation to generation. Molecular genetics employs the methods of genetics and molecular biology...
evidence is required to determine if the two taxa are genetically identical.
Ecology, habitat and distribution
Coprinellus micaceus is a saprotrophic species, deriving nutrients from dead and decomposing organic matterOrganic matter
Organic matter is matter that has come from a once-living organism; is capable of decay, or the product of decay; or is composed of organic compounds...
, and grows in and around stumps or logs of broad-leaved trees or attached to buried wood. It prefers feeding on bark
Bark
Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines and shrubs. Bark refers to all the tissues outside of the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner...
, particularly the secondary phloem, rather than the wood. In the scheme of the succession
Ecological succession
Ecological succession, is the phenomenon or process by which a community progressively transforms itself until a stable community is formed. It is a fundamental concept in ecology, and refers to more or less predictable and orderly changes in the composition or structure of an ecological community...
of fungal species involved in the decomposition of wood, C. micaceus is a late stage colonizer, and prefers to feed on wood that has already decomposed sufficiently to have reached "a friable softened consistency". A 2010 study suggests that the fungus can also live as an endophyte
Endophyte
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life without causing apparent disease. Endophytes are ubiquitous and have been found in all the species of plants studied to date; however, most of these endophyte/plant relationships...
, inhabiting the woody tissue of healthy trees without causing disease symptoms. The fungus is also associated with disturbed or developed ground, such as the sides of roads and paths, gardens, building sites and the edges of parking lots; it has also been noted for growing indoors on rotting wood in humid environments. In one instance it was discovered about 120 m (400 ft) underground in an abandoned coal mine, growing on wooden gangways and props used to support the roof.
Santiago de Compostela
Santiago de Compostela is the capital of the autonomous community of Galicia, Spain.The city's Cathedral is the destination today, as it has been throughout history, of the important 9th century medieval pilgrimage route, the Way of St. James...
in the Iberian Peninsula
Iberian Peninsula
The Iberian Peninsula , sometimes called Iberia, is located in the extreme southwest of Europe and includes the modern-day sovereign states of Spain, Portugal and Andorra, as well as the British Overseas Territory of Gibraltar...
, concluded that most "Coprinus" spores present in the atmosphere belonged to C. micaceus, and that the number of spores went up with increased humidity
Relative humidity
Relative humidity is a term used to describe the amount of water vapor in a mixture of air and water vapor. It is defined as the partial pressure of water vapor in the air-water mixture, given as a percentage of the saturated vapor pressure under those conditions...
and rainfall, but decreased with greater temperatures. The species is known for reappearing with successive fruitings at the same location. In one case, a total of 38 lb (17.2 kg) of fresh mushrooms were collected from one elm
Elm
Elms are deciduous and semi-deciduous trees comprising the genus Ulmus in the plant family Ulmaceae. The dozens of species are found in temperate and tropical-montane regions of North America and Eurasia, ranging southward into Indonesia. Elms are components of many kinds of natural forests...
stump in 10 successive crops over a spring and summer.
Coprinellus micaceus has a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...
, and has been collected in northern Africa, South Africa, Europe (including Turkey
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...
), North America (as far north as Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
), the Hawaiian islands, South America, India, Australia, New Zealand, and Japan. Phylogenetic analysis of rDNA
Ribosomal DNA
Ribosomal DNA codes for ribosomal RNA. The ribosome is an intracellular macromolecule that produces proteins or polypeptide chains. The ribosome itself consists of a composite of proteins and RNA. As shown in the figure, rDNA consists of a tandem repeat of a unit segment, an operon, composed of...
sequences from specimens collected in southeastern Asia and Hawaii show that the Hawaiian species form a distinct clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
with little genetic diversity
Genetic diversity
Genetic diversity, the level of biodiversity, refers to the total number of genetic characteristics in the genetic makeup of a species. It is distinguished from genetic variability, which describes the tendency of genetic characteristics to vary....
compared to Asian populations; this suggests that the Hawaiian populations have been introduced relatively recently and have not had much time to develop genetic variation. One study suggests that in South Africa, where C. micaceus is rare, it has been frequently confused with the similar-appearing C. truncorum
Coprinellus truncorum
Coprinellus truncorum is a species of mushroom in the Psathyrellaceae family. Part of the cluster of mushroom morphologically related to Coprinellus micaceus, this species can be distinguished from C. micaceus by a smooth, rather than pruinose stem, and by having more elliptical spores...
, a more common species in that region. A similar inference has been raised about North American species.
Bioactive compounds
Research into the natural productNatural product
A natural product is a chemical compound or substance produced by a living organism - found in nature that usually has a pharmacological or biological activity for use in pharmaceutical drug discovery and drug design...
chemistry of Coprinellus micaceus has revealed the presence of several chemical compounds unique to the species. Micaceol is a sterol
Sterol
Sterols, also known as steroid alcohols, are a subgroup of the steroids and an important class of organic molecules. They occur naturally in plants, animals, and fungi, with the most familiar type of animal sterol being cholesterol...
with "modest" antibacterial activity against the pathogens Corynebacterium xerosis and Staphylococcus aureus
Staphylococcus aureus
Staphylococcus aureus is a facultative anaerobic Gram-positive coccal bacterium. It is frequently found as part of the normal skin flora on the skin and nasal passages. It is estimated that 20% of the human population are long-term carriers of S. aureus. S. aureus is the most common species of...
. The compound (Z,Z)-4-oxo-2,5-heptadienedioic acid has inhibitory activity against glutathione S-transferase
Glutathione S-transferase
Enzymes of the glutathione S-transferase family are composed of many cytosolic, mitochondrial, and microsomal proteins. GSTs are present in eukaryotes and in prokaryotes, where they catalyze a variety of reactions and accept endogenous and xenobiotic substrates.GSTs can constitute up to 10% of...
, an enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
that has been implicated in the resistance of cancer cells against chemotherapeutic
Chemotherapy
Chemotherapy is the treatment of cancer with an antineoplastic drug or with a combination of such drugs into a standardized treatment regimen....
agents, especially alkylating
Alkylation
Alkylation is the transfer of an alkyl group from one molecule to another. The alkyl group may be transferred as an alkyl carbocation, a free radical, a carbanion or a carbene . Alkylating agents are widely used in chemistry because the alkyl group is probably the most common group encountered in...
drugs. A 2003 study did not find any antibacterial activity in this species. A 1962 publication reported the presence of the biologically active indole
Indole
Indole is an aromatic heterocyclic organic compound. It has a bicyclic structure, consisting of a six-membered benzene ring fused to a five-membered nitrogen-containing pyrrole ring. Indole is a popular component of fragrances and the precursor to many pharmaceuticals. Compounds that contain an...
compound tryptamine
Tryptamine
Tryptamine is a monoamine alkaloid found in plants, fungi, and animals. It is based around the indole ring structure, and is chemically related to the amino acid tryptophan, from which its name is derived...
in C. micaceus, although the concentration was not determined. The fruit bodies additionally produce a variety of pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...
compounds known as melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...
s—complex chemical polymer
Polymer
A polymer is a large molecule composed of repeating structural units. These subunits are typically connected by covalent chemical bonds...
s that contribute to the formation of soil humus after the fruit bodies have disintegrated. C. micaceus has been found to be devoid of the toxin coprine, the Antabuse-mimicking chemical found in Coprinus comatus
Coprinus comatus
Coprinus comatus, the shaggy ink cap, lawyer's wig, or shaggy mane, is a common fungus often seen growing on lawns, along gravel roads and waste areas. The young fruit bodies first appear as white cylinders emerging from the ground, then the bell-shaped caps open out. The caps are white, and...
that causes illness when consumed simultaneously with alcohol.