Cyathus stercoreus
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''Cyathus stercoreus'', commonly known as the dung-loving bird's nest or the dung bird's nest, is a species of fungus in the genus '' Cyathus'', family
Nidulariaceae The Nidulariaceae ('nidulus' - small nest) are a family of fungi in the order Agaricales. Commonly known as the bird's nest fungi, their fruiting bodies resemble tiny egg-filled birds' nests. As they are saprobic, feeding on decomposing organic ...
. Like other species in the Nidulariaceae, the fruiting bodies of ''C. stercoreus'' resemble tiny bird's nests filled with eggs. The fruiting bodies are referred to as splash cups, because they are developed to use the force of falling drops of water to dislodge and disperse their spores. The species has a worldwide distribution, and prefers growing on dung, or soil containing dung; the specific epithet is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''stercorarius'', meaning "of dung".Google Books
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Description

The fruiting bodies, or perida, are funnel- or barrel-shaped, 6–15 mm tall, 4–8 mm wide at the mouth, sometimes short-stalked, golden brown to blackish brown in age. The outside wall of the peridium, the ectoperidium, is covered with tufts of fungal hyphae that resembles shaggy, untidy hair. However, in older specimens this outer layer of hair (technically a ''tomentum'') may be completely worn off. The internal wall of the cup, the endoperidium, is smooth and grey to bluish-black. The 'eggs' of the bird's nest – the peridioles – are blackish, 1–2 mm in diameter, and there are typically about 20 in the cup. Peridioles are often attached to the fruiting body by a funiculus, a structure of hyphae that is differentiated into three regions: the basal piece, which attaches it to the inner wall of the peridium, the middle piece, and an upper sheath, called the purse, connected to the lower surface of the peridiole. In the purse and middle piece is a coiled thread of interwoven hyphae called the funicular cord, attached at one end to the peridiole and at the other end to an entangled mass of hyphae called the hapteron. However, Brodie reports that sometimes ''C. stercoreus'' is found without a funiculus, which has led some authors to misidentify this species with the genus '' Nidula''. The spores of ''C. stercoreus'' are roughly spherical and relatively large, with typical dimensions of 20–35 x 20–25  µm, although great variability in spore size has been noted.Brodie, ''The Bird's Nest Fungi'', p. 168. The spores are
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
(growing directly from the surface of the basidium, without attachment via a
sterigmata In biology, a sterigma (pl. sterigmata) is a small supporting structure. It commonly refers to an extension of the basidium (the spore-bearing cells) consisting of a basal filamentous part and a slender projection which carries a spore at the ti ...
), and are separated from the basidia after it collapses and gelatinizes. This is accompanied by the gelatinization of the inner walls of the peridiole.


Ultrastructure

Examination of fruiting bodies using scanning electron microscopy and
transmission electron microscopy Transmission electron microscopy (TEM) is a microscopy technique in which a beam of electrons is transmitted through a specimen to form an image. The specimen is most often an ultrathin section less than 100 nm thick or a suspension on a g ...
has revealed details about their ultrastructure—their microscopic architecture and arrangement. For example, the hyphae of the hapteron form a dense tangled network, while the hyphae of the funicular cord are arranged in a twisted form like a rope.Flegler SL, Hooper GR. (1978). "Ultrastructure of ''Cyathus stercoreus''".''Mycologia'' 70(6): 1181–1190. Further, the funicular cord, known to be highly elastic and with a high
tensile strength Ultimate tensile strength (UTS), often shortened to tensile strength (TS), ultimate strength, or F_\text within equations, is the maximum stress that a material can withstand while being stretched or pulled before breaking. In brittle materials t ...
, is made of thicker hyphae than the rest of the funiculus. Also, the ecto- and endoperidium are made of thick-walled, unbranched hyphae, known as skeletal hyphae. It has been proposed that these skeletal hyphae form a structural network that helps the fruiting body maintain the elasticity vital for proper functioning of the spore dispersal mechanism.


Life cycle

The life cycle of ''Cyathus stercoreus'', which contains both haploid and diploid stages, is typical of taxa in the
basidiomycetes Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Ba ...
that can reproduce both asexually (via vegetative spores), or sexually (with
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 r ...
).
Basidiospore A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are ...
s produced in the peridioles each contain a single haploid nucleus. After dispersal, the spores germinate and grow into homokaryotic hyphae, with a single nucleus in each compartment. When two homokaryotic hyphae of different mating compatibility groups fuse with one another, they form a
dikaryotic The dikaryon is a cell nucleus, nuclear feature which is unique to certain fungi. (The green alga ''Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types can fus ...
(containing two nuclei) mycelia in a process called
plasmogamy Plasmogamy is a stage in the sexual reproduction of fungi, in which the protoplasm of two parent cells (usually from the mycelia) fuse without the fusion of nuclei, effectively bringing two haploid nuclei close together in the same cell. This sta ...
. After a period of time (approximately 40 days when grown from
pure culture A microbiological culture, or microbial culture, is a method of multiplying microbial organisms by letting them reproduce in predetermined culture medium under controlled laboratory conditions. Microbial cultures are foundational and basic diagn ...
in the laboratory) and under the appropriate environmental conditions, fruiting bodies may be formed from the dikaryotic mycelia. These fruiting bodies produce peridioles containing the
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
upon which new basidiospores are made. Young basidia contain a pair of haploid sexually compatible nuclei which fuse, and the resulting diploid fusion nucleus undergoes meiosis to produce haploid basidiospores.


Development

Extreme variability in fruiting body form and color has been noted for ''C. stercoreus''. Brodie reported discovering a slender-stemmed "twinned" form, with two fruiting bodies originating from the same stalk. As has been shown in laboratory-grown specimens, the development and form of the fruiting bodies is at least partially dependent on the intensity of light it receives during development. For example, exposure of the heterokaryotic
mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...
to light is required for fruiting to occur, and furthermore, this light needs to be at a
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, t ...
of less than 530 nm. Lu suggests that certain growing conditions – such as a shortage in available nutrients – shifts the fungus'
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run c ...
to produce a hypothetical "photoreceptive precursor" that enables the growth of the fruiting bodies to be stimulated and affected by light. The fungi is also positively phototrophic, that is, it will orient its fruiting bodies in the direction of the light source.


Habitat and distribution

Being coprophilous, ''C. stercoreus'' grows on dung, in soil with dung, and bonfire sites; it has also been recorded growing on sand dunes. The fungus is known to have a worldwide distribution, and
Curtis Gates Lloyd Curtis Gates Lloyd (July 17, 1859 – November 11, 1926) was an American mycologist known for both his research on the gasteroid and polypore fungi, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with ab ...
, in his monograph on the Nidulariaceae, wrote that it "probably occurs in every country where manure occurs".


Spore dispersal

When a drop of water hits the interior of the cup at the appropriate angle and velocity, the peridioles are ejected into the air by the force of the drop. The force of ejection tears open the purse, and results in the expansion of the funicular cord, formerly coiled under pressure in the lower part of the purse. The peridioles, followed by the highly adhesive funicular cord and basal hapteron, may hit a nearby plant stem or stick. The hapteron sticks to it, and the funicular cord wraps around the stem or stick powered by the force of the still-moving peridiole. After drying out, the peridiole remains attached to the vegetation, where it may be eaten by a grazing herbivorous animal, and later deposited in that animal's dung to continue the life cycle.


Bioactive compounds

A number of
polyketide Polyketides are a class of natural products derived from a precursor molecule consisting of a chain of alternating ketone (or reduced forms of a ketone) and methylene groups: (-CO-CH2-). First studied in the early 20th century, discovery, biosynth ...
-type antioxidative compounds, cyathusals A, B, and C, and pulvinatal have been isolated and identified from the liquid culture of ''Cyathus stercoreus''. Furthermore, the polyketides known as cyathuscavin A, B, and C (isolated from liquid culture) also have antioxidant activity, and have DNA protection activity.


Uses

While inedible, the species has other uses.


Traditional medicine

In
Traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...
, a
decoction Decoction is a method of extraction by boiling herbal or plant material (which may include stems, roots, bark and rhizomes) to dissolve the chemicals of the material. It is the most common preparation method in various herbal-medicine systems. Dec ...
of this fungus is used to help relieve the symptoms of gastralgia, or stomach ache.


Agricultural and industrial

''Cyathus stercoreus'' has been investigated for its ability to break down lignin and
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
in agricultural byproducts, like
wheat Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeologi ...
straw or grasses. It selectively breaks down lignin, leaving much of the cellulose intact, which increases the amount of digestible
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or m ...
for
ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
mammals, and enhances both its value as a food source and its
biodegradability Biodegradation is the breakdown of organic matter by microorganisms, such as bacteria and fungi. It is generally assumed to be a natural process, which differentiates it from composting. Composting is a human-driven process in which biodegradati ...
. The enzymes responsible,
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 l ...
and
manganese peroxidase Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...
, also have industrial applications for lignin degradation and removal in the pulp and paper industry. Liquid cultures of ''C. stercoreus'' have also been shown to biodegrade the explosive compound 2,4,6-trinitrotoluene (
TNT Trinitrotoluene (), more commonly known as TNT, more specifically 2,4,6-trinitrotoluene, and by its preferred IUPAC name 2-methyl-1,3,5-trinitrobenzene, is a chemical compound with the formula C6H2(NO2)3CH3. TNT is occasionally used as a reagen ...
).Chen J. (1995). "Development of fungal degrading system to detoxify 2,4,6-trinitrotoluene (TNT) in liquid phase bioreactors". PhD dissertation, Texas A&M University. 127 pp.


See also

* List of ''Cyathus'' species


References


Cited text


External links

* {{Taxonbar, from=Q1265642 Nidulariaceae Medicinal fungi Fungi of Asia Fungi of Europe Fungi of North America Fungi of South America Fungi of Africa Fungi of Australia Fungi native to Australia Fungi described in 1888 Inedible fungi Taxa named by Lewis David de Schweinitz