Cyathus olla
Encyclopedia
Cyathus olla is a species of saprobic fungus in the genus Cyathus
Cyathus
Cyathus is a genus of fungi in the Nidulariaceae, a family collectively known as the bird's nest fungi. They are given this name since they resemble tiny bird's nests filled with "eggs", structures large enough to have been mistaken in the past for seeds. However, these are now known to be...

, family Nidulariaceae
Nidulariaceae
The Nidulariaceae are a family of fungi in the order Nidulariales. Commonly known as the bird's nest fungi, their fruiting bodies resemble tiny egg-filled birds' nests...

. The fruit bodies resemble tiny bird's nests filled with "eggs" – 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...

-containing structures called peridioles. Like other bird's nest fungi, C. olla relies on the force of falling water to dislodge peridioles from fruiting bodies to eject and disperse their spores. The life cycle
Biological life cycle
A life cycle is a period involving all different generations of a species succeeding each other through means of reproduction, whether through asexual reproduction or sexual reproduction...

 of this fungus allows it to reproduce both sexually, with 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....

, and asexually via 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...

s. C. olla is a relatively common fungus, with a worldwide distribution. It is the subject of agricultural research to determine its potential as a means to accelerate the breakdown of crop residue, and reduce the population of plant pathogens. The specific epithet is derived from the Latin word olla, meaning "pot".

Description

Cyathus olla bears a resemblance to a miniature bird's nest containing eggs, hence the common name bird's nest fungi. The fruiting body, or peridium, is roughly funnel-shaped, 10–18 mm tall and 8–12 mm wide. It is greyish yellow or greyish brown to fawn-colored and covered with even, fine-textured hairs on the outside surface. The inner surface is silver grey and smooth, often with faint transverse ridges. Cyathus olla is thick-walled, and flared outwards at the rim; the rim is typically wavy in outline. The 'eggs', or peridioles, typically number 8 to 10 in the cup, and they are white or grey in color, with a diameter of 2–4 mm—conspicuously larger than other Cyathus species. They are covered with a thin membrane called a tunica.
Peridioles are often attached to the fruiting body by a funiculus
Funiculus
The cerebrospinal fibers consist of numerous nerve fibers collected together and enclosed in membranous sheaths. A small bundle of fibers, enclosed in a tubular sheath, is called a funiculus; if the nerve is of small size, it may consist only of a single funiculus; but if large, the funiculi are...

, a structure of 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 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. Inside 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. C. olla has a wide purse with an upper portion not easily distinguishable from the lower part.

A different form of this species, Cyathus olla forma anglicus, originally reported from England by mycologist Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...

, has also been found in Oregon
Oregon
Oregon is a state in the Pacific Northwest region of the United States. It is located on the Pacific coast, with Washington to the north, California to the south, Nevada on the southeast and Idaho to the east. The Columbia and Snake rivers delineate much of Oregon's northern and eastern...

 and Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...

 in the USA, Alberta
Alberta
Alberta is a province of Canada. It had an estimated population of 3.7 million in 2010 making it the most populous of Canada's three prairie provinces...

 Canada, and in Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

. This form is large, with cup openings up to 1.5 cm (0.6 in) in diameter. Spores have dimensions of 11.5–12.5 × 7.5–9 µm.

The related species Cyathus earlei Lloyd closely resembles C. olla, although mating analysis
Mating in fungi
Mating in fungi is a complex process governed by mating types. Research on fungal mating has focused on only a few model species. Since not all of the fungi reproduce sexually and many that do are isogamous, the terms male and female do not apply to this kingdom...

 has shown that they are distinct species.

Microscopic features

Cyathus olla has ovate-shaped spores with dimensions of 10–14 × 6–8 µm
Micrometre
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...

. In 1927, G.W. Martin examined the microscopic features of various members of the Nidulariaceae family, including Cyathus olla. He noted that in this species, the basidia—spore-bearing cells—are club-shaped or cylindrical and have long stems, and they sometimes have a clamp connection
Clamp connection
A clamp connection is a structure formed by growing hyphal cells of certain fungi. It is created to ensure each septum, or segment of hypha separated by crossed walls, receives a set of differing nuclei, which are obtained through mating of hyphae of differing sexual types...

 at the basal end of the structure. Basidia typically hold between two to four sessile
Sessility (botany)
In botany, sessility is a characteristic of plants whose flowers or leaves are borne directly from the stem or peduncle, and thus lack a petiole or pedicel...

 spores that are directly attached to the basidium, rather than via a sterigma
Sterigma
thumb|right|The sterigmata is the slender extension that connects the spore to the basidia .A sterigma is an extension of the basidium consisting of a basal filamentous part and a slender projection with a spore at the tip...

. Revealed only after the spore is separated, a temporary apiculus marks the point of attachment. The spores are generally separated from the basidia when the latter collapse and gelatinize; this basidial breakdown typically occurs simultaneously with a gelatinization of cells lining the inside wall of the peridiole. After separating from the basidia the outer wall of the spores may thicken, although they generally do not increase in size.

Habitat and distribution

Being a saprobic fungus, Cyathus olla obtains nutrients from the breakdown of dead organic matter, and as such is usually found growing on woody debris; specimens found growing on soil are usually attached to bits of wood or stems present in the soil. Brodie notes that this species usually grows in moist and shaded locations, although its discovery in the arid regions of Lima
Lima
Lima is the capital and the largest city of Peru. It is located in the valleys of the Chillón, Rímac and Lurín rivers, in the central part of the country, on a desert coast overlooking the Pacific Ocean. Together with the seaport of Callao, it forms a contiguous urban area known as the Lima...

, Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....

, suggest that it is tolerant of low-moisture conditions. It is the most abundant species of Cyathus found in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, and is also common in North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

. Its range extends north to Sweden
Sweden
Sweden , officially the Kingdom of Sweden , is a Nordic country on the Scandinavian Peninsula in Northern Europe. Sweden borders with Norway and Finland and is connected to Denmark by a bridge-tunnel across the Öresund....

 and far south in South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

; it has also been reported in South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...

, Iran
Iran
Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...

, and Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

.

Life cycle

The life cycle of Cyathus olla, which contains both haploid and diploid stages, is typical of taxa in the basidiomycetes that can reproduce both asexually (via vegetative
Vegetative reproduction
Vegetative reproduction is a form of asexual reproduction in plants. It is a process by which new individuals arise without production of seeds or spores...

 spores), or sexually (with 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....

). Basidiospore
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...

s produced in the peridioles each contain a single haploid nucleus. After dispersal, the spores germinate and grow into homokaryotic
Homokaryotic
Homokaryotic refers to multinucleate cells where all nuclei are genetically identical. It is the antonym of heterokaryotic....

 hyphae, with a single nucleus in each compartment. When two homokaryotic 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 of different mating compatibility groups
Mating type
Mating types occur in eukaryotes that undergo sexual reproduction via isogamy. Since the gametes of different mating types look alike, they are often referred to by numbers, letters, or simply "+" and "-" instead of "male" and "female." Mating can only take place between different mating...

 fuse with one another, they form a dikaryotic
Dikaryon
Dikaryon is from Greek, di meaning 2 and karyon meaning nut, referring to the cell nucleus.The dikaryon is a nuclear feature which is unique to some fungi, in which after plasmogamy the two compatible nuclei of two cells pair off and cohabit without karyogamy within the cells of the hyphae,...

 mycelia in a process called plasmogamy
Plasmogamy
Plasmogamy is a stage in the sexual reproduction of fungi. In this stage, the cytoplasm of two parent mycelia fuse together without the fusion of nuclei, as occurs in higher terrestrial fungi. After plasmogamy occurs, the secondary mycelium forms. The secondary mycelium consists of dikaryotic...

. After a period of time and under the appropriate environmental conditions, fruiting bodies may be formed from the dikaryotic mycelia. These fruiting bodies 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 haploid basidiospores. Meiosis in C. olla has been found to be similar to that of higher organisms.

Spore dispersal

When a raindrop hits the interior of the cup with the proper angle and velocity it can produce considerable force, and can create a splash that drives the water up along the sides of the cup (also known as splash cups), tearing the funiculus, and ejecting the peridioles. The peridioles are followed by their funicular cord and basal hapteron. When they 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 (similar to a tetherball
Tetherball
Tetherball is a North American game for two opposing players. The equipment consists of a stationary metal pole, from which is hung a volleyball from a rope, or tether. The two players stand on opposite sides of the pole. Each player tries to hit the ball one way; one clockwise, and one...

). The peridiole, attached to the plant, may be eaten by herbivorous mammals, and the subsequent passage through its digestive tract will soften the hard shell enough to facilitate later sporulation.

Experiments investigating the splash dispersal of peridioles have shown a maximum horizontal distance for peridiole ejection of 82.5 centimetres (32.5 in), less than that observed for other Cyathus species. This reduced ejection distance may be due to such factors as the greater size of peridioles, the looser construction of the funiculus, or the increased flaring seen in the upper lip of the splash cup.

Agricultural applications

Blackleg is a fungal disease of canola
Canola
Canola refers to a cultivar of either Rapeseed or Field Mustard . Its seeds are used to produce edible oil suitable for consumption by humans and livestock. The oil is also suitable for use as biodiesel.Originally, Canola was bred naturally from rapeseed in Canada by Keith Downey and Baldur R...

 that results in major yield loss in affected crops. It overwinter
Overwinter
To overwinter is to pass through or wait out the winter season, or to pass through that period of the year when “winter” conditions make normal activity or even survival difficult or near impossible...

s on infected stubble (leftover crop residue) that is left on farm fields, and can continue to produce spores, infecting future crops, until the stubble is buried or broken down completely. The observation that C. olla grows and fruits on canola stubble has led to research on the potential of this fungus to degrade canola stubble, and reduce the incidence of stubble-borne diseases like blackleg and blackspot. In a study of its lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...

-degrading abilities, C. olla was shown to colonize canola, wheat
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

, and barley
Barley
Barley is a major cereal grain, a member of the grass family. It serves as a major animal fodder, as a base malt for beer and certain distilled beverages, and as a component of various health foods...

residue, but appeared to have a preference for the woody taproots of canola as compared to the cereal residue.

Cited text

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