Cyathus

Source: Wikipedia, the free encyclopedia.
Not to be confused with Cyathium or Cyathea.

Cyathus
Cyathus striatus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Agaricales
Family: Nidulariaceae
Genus: Cyathus
Haller (1768)
Type species
Cyathus striatus
(Huds.) Willd. (1787)
Species
Approximately 45[1]
Cyathus
saprotrophic
Edibility is inedible

Cyathus is a

endangered in a number of European countries. Some species of Cyathus are also known as splash cups, which refers to the fact that falling raindrops can knock the peridioles out of the open-cup fruit body. The internal and external surfaces of this cup may be ridged longitudinally (referred to as plicate or striate); this is one example of a taxonomic
characteristic that has traditionally served to distinguish between species.

Generally considered inedible, Cyathus species are

classification
systems that are based on traditional taxonomic characteristics.

Taxonomy

History

Bird's nest fungi were first mentioned by Flemish botanist

Harold J. Brodie in 1975.[9]

Infrageneric classification

The genus Cyathus was first subdivided into two infrageneric groups (i.e., grouping species below the rank of genus) by the Tulasne brothers; the "eucyathus" group had fruit bodies with inner surfaces folded into pleats (plications), while the "olla" group lacked plications.[5] Later (1906), Lloyd published a different concept of infrageneric grouping in Cyathus, describing five groups, two in the eucyathus group and five in the olla group.[7]

In the 1970s, Brodie, in his monograph on bird's nest fungi, separated the genus Cyathus into seven related groups based on a number of taxonomic characteristics, including the presence or absence of plications, the structure of the peridioles, the color of the fruit bodies, and the nature of the hairs on the outer peridium:[10]

  • Olla group: Species with a tomentum having fine flattened-down hairs, and no plications.
    • C. olla, C. africanus, C. badius, C. canna, C. colensoi, C. confusus, C. earlei, C. hookeri, C. microsporus, C. minimus, C. pygmaeus
  • Pallidus group: Species with conspicuous, long, downward-pointing hairs, and a smooth (non-plicate) inner peridium.
    • C. pallidus, C. julietae
  • Triplex group: Species with mostly dark-colored peridia, and a silvery white inner surface.
    • C. triplex, C. setosus, C. sinensis
  • Gracilis group: Species with tomentum hairs clumped into tufts or mounds.
    • C. gracilis, C. intermedius, C. crassimurus, C. elmeri
      The shaggy (tomentose) outer peridial surface of C. striatus
  • Stercoreus group: Species with non-plicate peridia, shaggy or wooly outer peridium walls, and dark to black peridioles.
    • C. stercoreus, C. pictus, C. fimicola
  • Poeppigii group: Species with plicate internal peridial walls, hairy to shaggy outer walls, dark to black peridioles, and large, roughly spherical or ellipsoidal spores.
    • C. poeppigii, C. crispus, C. limbatus, C. gayanus, C. costatus, C. cheliensis, C. olivaceo-brunneus
  • Striatus group: Species with plicate internal peridia, hairy to shaggy outer peridia, and mostly elliptical spores.
    • C. striatus, C. annulatus, C. berkeleyanus, C. bulleri, C. chevalieri, C. ellipsoideus, C. helenae, C. montagnei, C. nigro-albus, C. novae-zeelandiae, C. pullus, C. rudis

Phylogeny

The 2007 publication of

DNA sequence data of numerous Cyathus species has cast doubt on the validity of the morphology-based infrageneric classifications described by Brodie. This research suggests that Cyathus species can be grouped into three genetically related clades:[11]

C. dominicanus, an extinct species
  • Ollum group:
    • C. africanus (type), C. africanus f. latisporus, C. conlensoi, C. griseocarpus, C. guandishanensis, C. hookeri, C. jiayuguanensis, C. olla, C. olla f. anglicus, and C. olla f. brodiensis.
  • Striatum group:
  • Pallidum group:
    • C. berkeleyanus, C. olla f. lanatus, C. gansuensis, and C. pallidus.

This analysis shows that rather than fruit body structure, spore size is generally a more reliable character for segregating species groups in Cyathus.[11] For example, species in the ollum clade all have spore lengths less than 15 µm, while all members of the pallidum group have lengths greater than 15 µm; the striatum group, however, cannot be distinguished from the pallidum group by spore size alone. Two characteristics are most suited for distinguishing members of the ollum group from the pallidum group: the thickness of the hair layer on the peridium surface, and the outline of the fruit bodies. The tomentum of Pallidum species is thick, like felt, and typically aggregates into clumps of shaggy or woolly hair. Their crucible-shaped fruit bodies do not have a clearly differentiated stipe. The exoperidium of Ollum species, in comparison, has a thin tomentum of fine hairs; fruit bodies are funnel-shaped and have either a constricted base or a distinct stipe.[11]

Description

Species in the genus Cyathus have fruit bodies (peridia) that are vase-, trumpet- or urn-shaped with dimensions of 4–8 millimetres (3⁄16–5⁄16 inch) wide by 7–18 mm (1⁄4–11⁄16 in) tall.[12] Fruit bodies are brown to gray-brown in color, and covered with small hair-like structures on the outer surface. Some species, like C. striatus and C. setosus, have conspicuous bristles called setae on the rim of the cup. The fruit body is often expanded at the base into a solid rounded mass of hyphae called an emplacement, which typically becomes tangled and entwined with small fragments of the underlying growing surface, improving its stability and helping it from being knocked over by rain.[13]

Cyathus striatus (a) young and mature fruit bodies in longitudinal section; (b), (c) single peridiole entire, and in section

Immature fruit bodies have a whitish membrane, an epiphragm, that covers the peridium opening when young, but eventually dehisces, breaking open during maturation. Viewed with a microscope, the peridium of Cyathus species is made of three distinct layers—the endo-, meso-, and ectoperidium, referring to the inner, middle, and outer layers respectively. While the surface of the ectoperidium in Cyathus is usually hairy, the endoperidial surface is smooth, and depending on the species, may have longitudinal grooves (striations).[3]

Because the basic fruit body structure in all genera of the family Nidulariaceae is essentially similar, Cyathus may be readily confused with species of Nidula or Crucibulum, especially older, weathered specimens of Cyathus that may have the hairy ectoperidium worn off.[14] It distinguished from Nidula by the presence of a funiculus, a cord of hyphae attaching the peridiole to the endoperidium. Cyathus differs from genus Crucibulum by having a distinct three-layered wall and a more intricate funiculus.[3]

Peridiole structure

hapteron
  • g: middle piece

    • Derived from the Greek word peridion, meaning "small leather pouch",

    basidia, sterile (non-reproductive) structures, and spores. In young, freshly opened fruit bodies, the peridioles lie in a clear gelatinous substance which soon dries.[16]

    Peridioles are attached to the fruit body by a funiculus, a complex structure of hyphae that may be 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. In some species the peridioles may be covered by a tunica, a thin white membrane (particularly evident in C. striatus and C. crassimurus).[17] Spores typically have an elliptical or roughly spherical shape, and are thick-walled, hyaline or light yellow-brown in color, with dimensions of 5–15 by 5–8 µm.[12]

    Life cycle

    The life cycle of the genus Cyathus, which contains both

    mycelia, and the reproductive stage for the establishment of spore-producing structures, the fruit bodies.[18]

    The vegetative stage encompasses those phases of the life cycle involved with the germination, spread, and survival of the mycelium. Spores germinate under suitable conditions of moisture and temperature, and grow into branching filaments called hyphae, pushing out like roots into the rotting wood. These hyphae are

    saprobic, so mycelial growth in rotting wood is made possible by the secretion of enzymes that break down complex polysaccharides (such as cellulose and lignin) into simple sugars that can be used as nutrients.[19]

    After a period of time and under the appropriate environmental conditions, the dikaryotic mycelia may enter the reproductive stage of the life cycle. Fruit body formation is influenced by external factors such as season (which affects temperature and air humidity), nutrients and light. As fruit bodies develop they produce peridioles containing the basidia 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 basidiospores, each containing a single haploid nucleus.[20] The dikaryotic mycelia from which the fruit bodies are produced is long lasting, and will continue to produce successive generations of fruit bodies as long as the environmental conditions are favorable.

    Cyathus stercoreus

    The development of Cyathus fruit bodies has been studied in laboratory culture; C. stercoreus has been used most often for these studies due to the ease with which it may be grown experimentally.

    nm. Continuous light is not required for fruit body development; after the mycelium has reached a certain stage of maturity, only a brief exposure to light is necessary, and fruit bodies will form if even subsequently kept in the dark.[22] Lu suggested in 1965 that certain growing conditions—such as a shortage in available nutrients—shifts the fungus' metabolism to produce a hypothetical "photoreceptive precursor" that enables the growth of the fruit bodies to be stimulated and affected by light.[23] The fungi is also positively phototropic, that is, it will orient its fruit bodies in the direction of the light source.[24] The time required to develop fruit bodies depends on a number of factors, such as the temperature, or the availability and type of nutrients, but in general "most species that do fruit in laboratory culture do so best at about 25 °C, in from 18 to 40 days."[25]

    Bioactive compounds

    Structures of cyathin A3 (left) and cyathuscavins (right)[26][27]

    A number of species of Cyathus produce

    liquid culture of C. stercoreus, have significant antioxidant activity,[27] as do the compounds known as cyathusals, also from C. stercoreus.[35] Various sesquiterpene compounds have also been identified in C. bulleri, including cybrodol (derived from humulene),[36] nidulol, and bullerone.[37]

    Distribution and habitat

    Fruit bodies typically grow in clusters, and are found on dead or decaying wood, or on woody fragments in cow or horse dung.

    alpine plants at an altitude of 2,100 metres (7,000 ft).[42]

    C. poeppigii, a tropical species

    In general, species of Cyathus have a worldwide

    endangered in Bulgaria,[45] Denmark,[46] and Montenegro,[47] and "near threatened" in Great Britain.[48] The discovery of a Cyathus species in Dominican amber (C. dominicanus) suggests that the basic form of the bird's nest fungi had already evolved by the Cretaceous era and that the group had diversified by the mid-Cenozoic.[49]

    Ecology

    Spore dispersal

    Like other bird's nest fungi in the Nidulariaceae, species of Cyathus have their spores dispersed when water falls into the fruit body. The fruit body is shaped so that the kinetic energy of a fallen raindrop is redirected upward and slightly outward by the angle of the cup wall, which is consistently 70–75° with the horizontal.[50] The action ejects the peridioles out of the so-called "splash cup", where it may break and spread the spores within, or be eaten and dispersed by animals after passing through the digestive tract. This method of spore dispersal in the Nidulariaceae was tested experimentally by George Willard Martin in 1924,[51] and later elaborated by Arthur Henry Reginald Buller, who used C. striatus as the model species to experimentally investigate the phenomenon.[52] Buller's major conclusions about spore dispersal were later summarized by his graduate student Harold J. Brodie, with whom he conducted several of these splash cup experiments:

    Raindrops cause the peridioles of the Nidulariaceae to be thrown about four feet by splash action. In the genus Cyathus, as a peridiole is jerked out of its cup, the funiculus is torn and this makes possible the expansion of a mass of adhesive hyphae (the hapteron) which clings to any object in the line of flight. The momentum of the peridiole causes a long cord to be pulled out of a sheath attached to the peridiole. The peridiole is checked in flight and the jerk causes the funicular cord to become wound around stems or entangled among plant hairs. Thus the peridiole becomes attached to vegetation and may be eaten subsequently by herbivorous animals.[53]

    Although it has not been shown experimentally if the spores can survive the passage through an animal's digestive tract, the regular presence of Cyathus on cow or horse manure strongly suggest that this is true.[54] Alternatively, the hard outer casing of peridioles ejected from splash cups may simply disintegrate over time, eventually releasing the spores within.[55]

    Uses

    Species in the family Nidulariaceae, including Cyathus, are considered inedible, as (in Brodie's words) they are "not sufficiently large, fleshy, or odorous to be of interest to humans as food".

    human physiology is unknown.[57]

    Biodegradation

    Cyathus olla; note the smooth (not plicate) endoperidium, and relatively large peridioles.

    munitions-contaminated soils.[62]

    Agriculture

    mineralization of essential plant nutrients.[63][64]

    Human biology

    Various Cyathus species have antifungal activity against human pathogens such as Aspergillus fumigatus, Candida albicans and Cryptococcus neoformans.[65] Extracts of C. striatus have inhibitory effects on NF-κB, a transcription factor responsible for regulating the expression of several genes involved in the immune system, inflammation, and cell death.[66]

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    Cited texts

    External links

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