Basidium
Encyclopedia
A basidium is a microscopic, spore-producing structure found on the hymenophore
Hymenophore
A hymenophore refers to the hymenium-bearing structure of a fungal fruiting body. Hymenophores can be smooth surfaces, lamellae, folds, tubes, or teeth....

 of fruiting bodies of basidiomycete fungi. The presence of basidia is one of the main characteristic features of the Basidiomycota
Basidiomycota
Basidiomycota is one of two large phyla that, together with the Ascomycota, comprise the subkingdom Dikarya within the Kingdom Fungi...

. A basidium usually bears four sexual spores called basidiospores; occasionally the number may be two or even eight. In a typical basidium, each basidiospore is borne at the tip of a narrow prong or horn called 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...

 (pl. sterigmata), and is forcibly discharged upon maturity.

The word basidium literally means little pedestal, from the way in which the basidium supports the spores. However, some biologists suggest that the structure more closely resembles a club
Club (weapon)
A club is among the simplest of all weapons. A club is essentially a short staff, or stick, usually made of wood, and wielded as a weapon since prehistoric times....

. An immature basidium is known as a basidiole.

Basidium structure

Most basidiomycetes have single celled basidia (holobasidia), but in some groups basidia can be multicellular (a phragmobasidia). For instance, rust fungi
Rust (fungus)
Rusts are plant diseases caused by pathogenic fungi of the order Pucciniales. About 7800 species are known. Rusts can affect a variety of plants; leaves, stems, fruits and seeds. Rust is most commonly seen as coloured powder, composed off tiny aeciospores which land on vegetation producing...

 in the order Uredinales have four-celled phragmobasidia that are transversely septate; some jelly fungi in the order Tremellales
Tremellales
The Tremellales are an order of fungi in the class Tremellomycetes. The order contains both teleomorphic and anamorphic species, most of the latter being yeasts. All teleomorphic species in the Tremellales are parasites of other fungi, though the yeast states are widespread and not restricted to...

 have four-celled phragmobasidia that are cruciately septate. Sometimes the basidium (metabasidium) develops from a probasidium, which is a specialized cell which is not elongated like a typical 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...

. The basidium may be stalked or sessile.

Mechanism of basidiospore discharge

In most basidiomycetes, the basidiospores are ballistospores--they are forcibly discharged. The propulsive force is derived from a sudden change in the center of gravity
Center of gravity
In physics, a center of gravity of a material body is a point that may be used for a summary description of gravitational interactions. In a uniform gravitational field, the center of mass serves as the center of gravity...

 of the discharged spore. Important factors in forcible discharge include Buller's drop, a droplet of fluid that can be observed to accumulate at the proximal tip (hilar appendage) of each basidiospore; the offset attachment of the spore to the subtending sterigma, and the presence of hygroscopic regions on the basidiospore surface.

Upon maturity of a basidiospore, sugars present in the cell wall begin to serve as condensation loci for water vapor in the air. Two separate regions of condensation are critical. At the pointed tip of the spore (the hilum) closest to the supporting basidium, Buller's drop accumulates as a large, almost spherical water droplet. At the same time, condensation occurs in thin film on the adaxial face of the spore. When these two bodies of water coalesce, the release of surface tension and the sudden change in the center of mass leads to sudden discharge of the basidiospore. Remarkably, Money (1998) has estimated the initial acceleration of the spore to be about 10,000 g.

Successful basidiospore discharge can only occur when there is sufficient water vapor available to condense on the spore.

Evolutionary loss of forcible discharge

Some basidiomycetes lack forcible discharge, although they still form basidiospores. In each of these groups, spore dispersal occurs through other discharge mechanisms. For example, members of the order Phallales (stinkhorns) rely on insect vectors for dispersal; the dry spores of the Lycoperdales
Lycoperdales
The Lycoperdales are a now outdated order of fungi. The order included some well-known types such as the giant puffball, the earthstars, and other tuberous fungi...

 (puffball
Puffball
A puffball is a member of any of several groups of fungus in the division Basidiomycota. The puffballs were previously treated as a taxonomic group called the Gasteromycetes or Gasteromycetidae, but they are now known to be a polyphyletic assemblage. The distinguishing feature of all puffballs is...

s) and Sclerodermataceae
Sclerodermataceae
The Sclerodermataceae are a family of fungi in the order Boletales, containing several genera of unusual fungi that little resemble boletes. Taxa, which include species commonly known as the ‘hard-skinned puffballs’, ‘earthballs’, or 'earthstars', are widespread in both temperate and tropical regions...

 (earth balls and kin) are dispersed when the basidiocarps are disturbed; and species of the Nidulariales (bird's nest fungi) use a splash cup mechanism. In these cases the basidiospore typically lacks a hilar appendage, and no forcible discharge occurs. Each example is thought to represent an independent evolutionary loss of the forcible discharge mechanism ancestral to all basidiomycetes.

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