Coprinellus micaceus - Wikipedia https://en.wikipedia.

org/wiki/Coprinellus_micaceus

Coprinellus micaceus
Coprinellus micaceus is a common species of fungus in the
family Psathyrellaceae with a cosmopolitan distribution. The fruit
Coprinellus micaceus
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 or
linear 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
Scientific classification
the common names mica cap, shiny cap, and glistening inky
cap. Although small and with thin flesh, the mushrooms are usually Kingdom: Fungi
bountiful, as they typically grow in dense clusters. A few hours after Division: Basidiomycota
collection, the gills will begin to slowly dissolve into a black, inky,
Class: Agaricomycetes
spore-laden liquid—an enzymatic process called autodigestion or
deliquescence. The fruit bodies are edible before the gills blacken and Order: Agaricales
dissolve, and cooking will stop the autodigestion process. Family: Psathyrellaceae
The microscopic characteristics and cytogenetics of C. micaceus are Genus: Coprinellus
well known, and it has been used frequently as a model organism to Species: C. micaceus
study cell division and meiosis in Basidiomycetes. Chemical analysis
of the fruit bodies has revealed the presence of antibacterial and
Binomial name
enzyme-inhibiting compounds. Formerly known as Coprinus Coprinellus micaceus
micaceus, the species was transferred to Coprinellus in 2001 as (Bull.:Fr.) Vilgalys, Hopple & Jacq.
phylogenetic analyses provided the impetus for a reorganization of Johnson
the many species formerly grouped together in the genus Coprinus.
Synonyms[1]
Based on external appearance, C. micaceus is virtually
indistinguishable from C. truncorum, and it has been suggested that Agaricus micaceus Bull.
many reported collections of the former may be of the latter. Coprinus micaceus (Bull.) Fr.

Coprinellus micaceus
Contents Mycological characteristics
History and taxonomy
gills on hymenium
Description
Microscopic characteristics
cap is campanulate or
Edibility
convex
Similar species
Ecology, habitat and distribution hymenium is adnexed

Bioactive compounds
stipe is bare
See also
References spore print is black

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Cited books ecology is

saprotrophic
External links
edibility: edible

History and taxonomy

Agaricus micaceus, illustrated by
Bulliard in 1786
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.[9]
Coprinellus micaeus was illustrated in a woodcut 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]).[2]
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.[3] 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.[4] 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.[5] 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 hyphae intermixed.[6]
Molecular studies published in the 1990s[7][8] demonstrated that many

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,[10] 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.[2] 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.[11]

The specific epithet micaceus is derived from the Latin word mica, for "crumb, grain of salt" and the suffix -aceus,
"like, similar";[12] the modern application of "mica" to a very different substance comes from the influence of
micare, "glitter".[13] The mushroom is commonly known as the "shiny cap",[14] the "mica cap" or the "glistening
inky cap", all in reference to the mealy particles found on the cap that glisten like mica.[15]

Description
The cap is initially 1–2.5 cm (0.4–1.0 in) 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

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convex. When expanded, the cap diameter reaches 0.8–3.0 cm

(0.3–1.2 in) 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
Gills of young... veil that covers immature specimens.[16] The particles are loosely attached
and easily washed away, so that older specimens are often smooth.[17]
Coprinellus micaceus is hygrophanous, meaning it assumes different
colors depending on its state of hydration.[18]

The gills are crowded together closely, and have an adnexed (narrow)
attachment to the stem.[19] Initially white, they change color to dark
brown then eventually black as the spores mature. Expansion of the cap
...and old fruit bodies 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.[20] The brittle stem is hollow, and measures 4–10 cm (1.6–3.9 in) long by 0.2–0.5 cm
(0.1–0.2 in) 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.[15] The spore print is dark brown or black.[21] The flesh is thin, fragile, white
in the stem, and brownish in the cap.[22] Its odor and taste are not distinctive.[23] Individual fruit bodies take an
average of five to seven days to fully mature.[24]

Microscopic characteristics
The spores of C. micaceus are reddish-brown, with dimensions of 7–10
by 4.5–6 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,
a flattened area in the center of the spore surface through which a germ
tube may emerge.[21] The spore-bearing cells (the basidia) are four-
spored, club-shaped, and measure 10–15 by 4–7 µm.[25] Studies have
Spores are mitriform (shield-
shown that the basidia develop in four discrete generations. The first shaped)
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.[26]

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

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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.[27] 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.[25] 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,[28]
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 vacuoles 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.[29]

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.[20] 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."[30]

In 1914, Michael Levine was the first to report successfully cultivating C. 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.[31] Like other coprinoid species, C. micaceus
undergoes synchronous meiosis. The chromosomes 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.[32][33] The chromosome number of C. micaceus is n=12.[34]

Edibility
Coprinellus micaceus is an edible species,[19][35] and cooking inactivates the enzymes that cause autodigestion or
deliquescence—a process that can begin as soon as one hour after collection.[36] It is considered ideal for
omelettes,[22] and as a flavor for sauces,[14] although it is "a very delicate species easily spoiled by overcooking".[37]
The flavor is so delicate that it is easy to overpower and hide with almost anything. The fungus also appeals to fruit
flies of the genus Drosophila, who frequently use the fruit bodies as hosts for larvae production.[38][39]

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.[40] 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.[41]

Similar species
The edible Coprinellus bisporus 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).[42] 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.[43] Coprinellus truncorum is also covered with glistening granules

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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.[44] 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.[25]

Ecology, habitat and distribution

Coprinellus micaceus is a saprotrophic species, deriving nutrients from
dead and decomposing organic matter, and grows in and around
stumps or logs of broad-leaved trees or attached to buried wood. It
The related species Coprinellus
prefers feeding on bark, particularly the secondary phloem, rather than
disseminatus invariably grows in
the wood.[45] In the scheme of the succession of fungal species involved large clusters on wood.
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".[46] A 2010 study suggests that the fungus can also live as an
endophyte, inhabiting the woody tissue of healthy trees without causing disease symptoms.[47] 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;[48] it has also been noted for growing indoors on rotting wood in humid
environments.[15] 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.[49]

Fruit bodies are commonly found growing in dense clusters, but can
also be found growing singly or in small clumps, especially in forested
areas.[18] In North America, C. micaceus is one of the first edible
mushrooms to appear in the spring,[23] and fruits from April to
September. In Europe, it fruits from May to December.[37] 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.[43] A study of air quality conducted in the city of
A cluster of fruit bodies at the base
Santiago de Compostela in the Iberian Peninsula, concluded that most
of a tree in Wayne National Forest,
"Coprinus" spores present in the atmosphere belonged to C. micaceus,
Ohio, USA
and that the number of spores went up with increased humidity and
rainfall, but decreased with greater temperatures.[50] 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.[51][52]

Coprinellus micaceus has a cosmopolitan distribution,[22] and has been collected in northern Africa,[53] South
Africa,[54] Europe (including Turkey[55]), North America (as far north as Alaska),[56] the Hawaiian islands,[25][44]
South America,[25] India,[57][58] Australia,[37] New Zealand,[59] and Japan.[60] 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.[61]
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.[54] A similar inference has been raised

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about North American species.[56][62]

Bioactive compounds
Research into the natural product 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.[63][64] A 2003 study did not find any
antibacterial activity in this species.[65] A 1962 publication reported the presence of the biologically active indole
compound tryptamine in C. micaceus, although the concentration was not determined.[66] The fruit bodies
additionally produce a variety of pigment compounds known as melanins—complex chemical polymers that
contribute to the formation of soil humus after the fruit bodies have disintegrated.[67] C. micaceus has been found
to be devoid of the toxin coprine, the Antabuse-mimicking chemical found in Coprinopsis atramentaria that
causes illness when consumed simultaneously with alcohol.[68]

See also
List of Coprinellus species

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Cited books
Arora D. (1986). Mushrooms Demystified: a Comprehensive Guide to the Fleshy Fungi. Berkeley, California:
Ten Speed Press. ISBN 0-89815-169-4.
Buller AH (1924). "Chapter XI: The micaceus sub-type illustrated by Coprinus micaceus". Researches on
Fungi. III. London: Longmans, Green, and Co. pp. 328–56.

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