Introduction

The Basidiomycota contains about 30,000 described species, which is 37% of the described species of true Fungi (Kirk et al. 2001). The most conspicuous and familiar Basidiomycota are those that produce mushrooms, which are sexual reproductive structures. The Basidiomycota also includes yeasts (singlecelled forms; Fell et al. 2001) and asexual species. Basidiomycota are found in virtually all terrestrial ecosystems, as well as freshwater and marine habitats (Kohlmeyer and Kohlmeyer, 1979; Hibbett and Binder, 2001). Basidiomycota have a huge impact on human affairs and ecosystem functioning. Many Basidiomycota obtain nutrition by decaying dead organic matter, including wood and leaf litter. Thus, Basidiomycota play a significant role in the carbon cycle. Unfortunately, Basidiomycota frequently attack the wood in buildings and other structures, which has negative economic consequences for humans. Symbiotic lifestyles (intimate associations with other living organisms) are well developed in the Basidiomycota. Symbiotic Basidiomycota include important plant pathogens, such as "rusts" (Uredinales) and "smuts" (Ustilaginales), which attack wheat and other crops. Other symbiotic Basidiomycota cause diseases in animals, including humans. Not all symbiotic Basidiomycota cause obvious harm to their partners, however. For example, some Basidiomycota, as well as a handful of Ascomycota, form ectomycorrhizae, which are associations with the roots of vascular plants (principally forest trees such as oaks, pines, dipterocarps, and eucalypts; Smith and Read, 1997). Ectomycorrhizal Basidiomycota help their plant partners obtain mineral nutrients from the soil, and in return they receive sugars that the plants produce through photosynthesis. Other symbiotic Basidiomycota form associations with insects, including leaf-cutter ants, termites, scale insects, woodwasps, and bark beetles (Wheeler and Blackwell, 1984; Mueller et al., 1998). Humans have found diverse uses for Basidiomycota. Mushrooms, both cultivated and wild, are eaten in many countries. For the untrained, mushroom-hunting is a risky endeavor, because some Basidiomycota produce deadly toxins (Benjamin 1995). The basidiomycete toxin phalloidin (from the mushroom Amanita phalloides) binds actin, which is a component of microfilaments. Fluorescent stains that incorporate phalloidin are used by cell biologists to visualize the cytoskeleton. Other "toxins" produced by Basidiomycota include hallucinogens, which are produced by members of the genus Psilocybe (and other groups). Species of Psilocybe have traditionally been used in Central American indigenous cultures as a spiritual tool, and are now cultivated for the illicit drug trade. Other biochemical compounds of Basidiomycota that have practical uses include astaxanthin, a red pigment produced by the basidiomycetous yeast Phaffia (used to add color to farmed salmon), and certain enzymes from wood-decaying Basidiomycota that have potential applications in paper production and bioremediation (decontamination of polluted environments using biological agents).

Characteristics
Basidiomycota are unicellular or multicellular, sexual or asexual, and terrestrial or aquatic. Indeed, Basidiomycota are so variable that it is impossible to identify any morphological characteristics that are both unique to the group and constant in the group. The most diagnostic feature is the production of basidia (sing. basidium), which are the cells on which sexual spores are produced, and from which the group takes its name. A long-lived dikaryon, in which each cell in the thallus contains two haploid nuclei resulting from a mating event, is another characteristic feature. Finally, clamp connections are a kind of hyphal outgrowth that is unique to Basidiomycota, although they are not present in all Basidiomycota. The following description of the characteristics of Basidiomycota traces the life cycle of a "typical" species, beginning at the site of meiosis. The basidium is the cell in which karyogamy (nuclear fusion) and meiosis occur, and on which haploid basidiospores are formed (basidia are not produced by asexual Basidiomycota). Many Basidiomycota produce basidia on multicellular fruiting bodies (e.g., mushrooms), but basidia can also be formed directly from yeasts or other single cells. There is a great range of variation in morphology of the basidium, the number of spores formed, and how the spores are borne on the surface of the basidium (Ingold 1991). Typically, four spores are produced on each basidium, at the tips of minute stalks called sterigmata (Fig. 1). Each spore usually contains one or two of the haploid meiotic products.

Figure 1. SEM of the surface of a mushroom gill (Coprinus cinereus: Agaricomycotina) showing several basidia, some with four basidiospores attached. (From McLaughlin, et al. 1985; used with permission McLaughlin, Beckett and Yoon 1985.)

One of the most fascinating characteristics of Basidiomycota is the production of forcibly discharged ballistospores (Fig. 2), which are propelled into the air from the sterigma. Ballistospores may be sexual or asexual, and may be produced by basidia, hyphae, yeast cells, or even other ballistospores. This type of spore discharge must have evolved very early in the evolutionary history of the Basidiomycota as it is found in members of the earliest diverging lineages within the group. Ballistospory is associated with forms that disperse their spores directly into the air. Most aquatic Basidiomycota and forms that produce spores inside the fruiting body, such as puffballs, have lost ballistospory. Ballistospory is associated with the production of a liquid filled "hilar droplet" that forms at the base of the spore, just above its attachment to the sterigma (Fig. 2). Resolving the mechanism of ballistospory has been a longstanding problem in mycology (Buller 1909, 1922; Ingold 1939; McLaughlin et al. 1985; Webster et al. 1984a, b; Yoon and McLaughlin 1986). In a series of studies, reviewed by Money (1998), it has been shown that spore discharge occurs when the hilar droplet fuses with a film of liquid on the surface of the spore. The rapid coalescence of the liquids causes a sudden shift in the center of mass of the spore and contributes to its release from the sterigma. This mechanism has been termed a "surface tension catapult" and it results in spores being discharged with a force of about 25,000 g (Money 1998, Pringle et al. 2005).

Figure 2. Scanning electron micrograph showing the hilar droplet at the base of a basidiospore (Coprinus cinereus: Agaricomycotina). (From McLaughlin et al. 1985)

Basidiospores germinate to form hyphae (filaments) or yeast cells that are typically haploid and uninucleate. The hyphae of Basidiomycota are septate. Ultrastructural features of the septa, visible with transmission electron microscopy, have been important in developing phylogenetic hypotheses in Basidiomycota (see theAgaricomycotina page). Mating in Basidiomycota involves fusion of haploid cells, but fusion of the nuclei is usually delayed until the basidia are formed. Thus, the dominant phase of the life cycle in most Basidiomycota is a dikaryon, in which the two nuclei brought together in mating exist side-by-side in each cell (Fig. 3A). Sometimes a dikaryon can donate a nucleus to a uninucleate "monokaryon", resulting in a "di-mon" mating. Clamp connections are hyphal outgrowths that form when cells in dikaryotic hyphae divide. One of the nuclei divides in the main axis of the hypha, while the other divides into the clamp (Fig. 3B). Septa are formed across each of the mitotic spindles. The apex of the backward-growing monokaryotic clamp cell fuses with the subapical cell, reestablishing the dikaryotic condition (Fig. 3C). All fungi that produce clamp connections are members of the Basidiomycota, but not all Basidiomycota produce clamp connections. The regular formation of clamp connections must have developed early in basidiomycete evolution, because they are found in all the major clades of Basidiomycota. Sexually reproducing Ascomycota also form dikaryons, although they are not as long-lived as those of Basidiomycota. The clade that includes Ascomycota and Basidiomycota has been called the Dikaryomycotina, reflecting this presumably homologous similarity. Ascomycota produce clamp-like "croziers" at the bases of asci (cells in which meiosis occurs, homologous to basidia). Croziers may be homologous to clamp connections.

Figure 3. Diagram of clamp cell formation: A. Dikaryotic hypha, arrow shows direction of hyphal tip growth. B. Clamp cell growing backward, nuclei undergoing synchronous division. C. Mature clamp. Illustrations E. Swann 1997.

Discussion of Phylogenetic Relationships

There is strong evidence that the Basidiomycota is monophyletic. Ballistospores, basidia, and clamp connections are present in the Agaricomycotina, Ustilaginomycotina, and Pucciniomycotina (although not in all species), suggesting that they have a common origin. Non-molecular characters that have been used to recognize major groups within the Basidiomycota include the form of the basidia (shape and septation), ultrastructure of hyphal septa and spindle pole bodies, presence or absence of yeast phases and "spore repetition" (production of spores directly from spores), and cellular carbohydrate composition (McLaughlin et al. 1995; Oberwinkler, 1987; Prillinger et al. 1990, 1991). Sequences of ribosomal RNA (rRNA) genes, recently supplemented by protein-coding genes, have played a major role in increasing our understanding of the relationships within Basidiomycota, and have demonstrated that some morphological attributes that have been emphasized in higher-level classification, such as the form of basidia, are subject to homoplasy (Swann and Taylor 1993, 1995, Swann et al. 1999). Three major clades are strongly supported within the Basidiomycota: 1) Pucciniomycotina includes rusts (Pucciniales) and other taxa (Swann et al. 2001, Aime et al. 2006); 2) Ustilaginomycotina includes smuts (Ustilaginales) and others (Bauer et al. 2001, Begerow et al. 2006); and 3) Agaricomycotina includes mushrooms (Agaricomycetes), jelly fungi (Auriculariales, Dacrymycetales, Tremellales) and others (Hibbett and Thorn 2001, Swann and Taylor 1995, Wells and Bandoni 2001, Hibbett 2006).

Monophyly of each of these groups has been supported in phylogenetic analyses of rRNA gene sequences and protein-coding genes (Hibbett et al. 2007). Similarities in the ultrastructure of septal pores and spindle pole bodies (McLaughlin et al. 1995) suggest that Ustilaginomycotina and Agaricomycotina could be sister groups, and some molecular phylogenies also support this topology. The placements of the Wallemiomycetes (a group of osmophilic molds) and Entorrhizomycetes (a group of root-inhabiting Fungi, previously classified in the Ustilaginomycotina [Bauer et al. 2001]) are particularly problematical (Matheny et al. 2006). At present, these are classified as "incertae sedis" within the Basidiomycota (Hibbett et al. 2007), but with the application of genome-scale datasets their placements may be resolved.

Introduction
The fungal group basidiomycota is best known for the production of large fruitbodies such as the mushrooms, puffballs, brackets, etc. However, the group also contains some microscopic fungi, including the important rust fungi and smut fungi that parasitise plants (see Biotrophic parasites), and some yeasts. One of these yeasts, Sporobolomyces roseus, is very common on moribund leaf surfaces and is a cause for concern because its basidiospores are respiratory allergens. Another is Cryptococcus neoformans, which grows commonly on old "weathered" bird droppings, and which can cause a fatal systemic infection of immunocompromised people. Its air-borne basidiospores initiate infection via the lungs, leading to the disease termed cryptococcosis. This is now the fourth most important lifethreatening diseases of AIDS patients in the USA, and common also in Europe. Many of the basidiomycota with the larger fruitbodies (toadstools etc.) are common and important agents of wood decay (see Wood decay) or decomposers of leaf litter, animal dung, etc. Others are mycorrhizal on forest trees (see Mycorrhizas). Several species are grown commercially for food, including the common cultivated mushroom, Agaricus bisporus, and some more exotic species which can be found on supermarket shelves. For example, the supermarket package below contains the brown-coloured Shiitake mushroom (Lentinus edodes, traditionally grown in south-east Asia) and a mixture of grey, pink and yellow forms of the oyster fungus (Pleurotus ostreatus, reportedly an aphrodisiac).

Distinctive features and life cycle of the basidiomycota

As a group, the basidiomycota have some highly characteristic features, which separate them from other fungi. They are the most evolutionarily advanced fungi, and even their hyphae have a dinstinctly "cellular" composition. This point is best illustrated by the life cycle below.

The basidiospores usually have a single haploid nucleus. When these spores germinate they produce hyphae with usually a single nucleus in each compartment. Because these hyphae have only one nuclear type they are termed monokaryons (from the Greek, mono = one; karyos = kernel or nucleus). At some stage in their growth, two monokarons of different compatibility groups (mating types) fuse. This can occur either by simple hyphal fusions or by fusion of a hypha with a small spore termed an oidium. The fusion event is termed plasmogamy. Then the nuclei divide and the daughter nuclei pair so that each hyphal compatment comes to contain two nuclei - one of each mating type. At this stage the fungus is termed a dikaryon (i.e. with two nuclear types). Many basidiomycota grow for most of their lives as dikaryons, until environmental signals induce them to produce fruibodies, such as a toadstool. All the hyphae that make up the toadstool are dikaryotic. At a late stage of development, some of these hyphae produce special cells termed basidia (singular, basidium). For example, the cells that line the gills of the common mushroom are basidia. Finally, the two haploid nuclei in each basidium fuse - a process termed karyogamy) to form a diploid nucleus. This then undergoes meiosis to produce four haploid nuclei, and these haploid nuclei migrate into the basidiospores, which develop on small stalks (termed sterigmata) from each basidium (see the image below). The dispersal of these monokaryotic spores completes the life cycle.

Scanning electron micrograph of basidia on the gills of a toadstool. Note that each basidium produces four stalks (sterigmata) and the basidiospores develop on the ends of these stalks. [Image courtesy of Dr C. Jeffree] In many basidiomycota there is a rather elaborate mechanism for ensuring that the dikaryotic condition is maintained during growth of the hyphae. As shown in the diagram below, the two nuclei in each tip cell divide at the same time, but one divides along the axis of the hypha, and the other divides so that a daughter nucleus enters a small, backwards-projecting branch. Then a septum forms to separate the original apical cell into two cells, and the branch fuses with the sub-terminal cell so that the nucleus in the branch migrates into this cell. The small branches at each septum are termed clamp connections. They can be seen at high magnification of a normal compound microscope (see the image below the diagram).

Diagram to show the role of clamp connections in maintaining a dikaryon [From JW Deacon, 1997, Modern Mycology, Blackwell Science]

Clamp connections at the septa of a basidiomycota hypha

Some unusual fruitbodies

Many basidiomycota produce a typical toadstool-shaped fruitbody, with basidia lining the gills (or sometimes pores instead of gills). But an assemblage of basidiomycota termed gasteromycetes produce some unusual types of fruitbody, with basidia that mature inside the fruitbody. Puffballs (see below) are common examples of this - the basidiospores form a powdery mass inside the fruitbody. At maturity, the outer layer of the puffball becomes a papery structure with a hole at the top, and a "puff" of basidiospores is released when raindrops impact onto the fruibody.

Immature puffballs developing on rotten wood. Note the thick white strands on the surface of the wood. Each of these "mycelial strands" is composed of many hyphae, which channel nutrients to the developing fruitbodies from the mycelium growing in the wood.

Earthstars (above and below) are the fruitbodies of Geastrum and similar genera. They are like puffballs, but often develop from mycelium growing below ground in woodlands. In order to ensure that the fruitbodies are not covered in leaf litter, the outer layers of the fruitbody split and fold back at maturity, raising the inner "puffball" above ground level.

Five fruitbodies of Geastrum, in various stages of "unfolding" The "bird's nest fungi (see below) commonly grow on wood chippings. Their fruitbodies consist of a splash-cup, about 1 cm diameter. Within this are clusters of "eggs", termed peridioles. These are forcibly ejected when raindrops fall into the splash-cup, and they land on the surrounding ssubstrate, where they adhere by means of a sticky pad (termed a hapteron).

Cyathus (bird's nest) fruitbodies developing from hyphae growing in wood chips. Note the funnel-shaped fruitbodies and the clusters of "eggs" inside them. The same features are shown in the image below, after a rainstorm, when some of the peridioles (eggs) were splashed onto the surrounding wood (arrowheads). The peridiole indicated by the arrowhead near the top centre of this image is seen to have a sticky attachment on its right side.

Fairy ring fungi

Several basidiomycota grow as decomposers on the "thatch" of dead leaves and roots that accumulate in old grasslands. Their produce rings of fruitbodies in autumn, known as fairy rings because folklore associates them

with the circles of fairy dances. The rings expand each year, as the fungal hyphae grow into new zones of thatch, and the hyphae in the older zone die, releasing nitrogen into the soil. This nitrogen promotes the growth of grass behind the advancing margin of the ring, giving it a richer green colour. Thus, we can see fairy rings for much of the year, as rings of enhanced grass growth, even when the fungal fruitbodies are not present. Some fairy rings are extremely large - for example those on the grass plains around Stonehenge in Wiltshire - and measurements of their current annual rates of expansion suggest that such rings can be at least 400 years old. Most fairy ring fungi produce only "green" rings and cause no damage to the grass. For example, this is true of the rings caused by Hygrocybe species, the fruitbodies of which are waxy and often brightly coloured.

Rings of Hygrocybe fruitbodies and zones of lush grass growth caused by this fungus.

Fruitbodies of Hygrocybe from non-killing fairy rings However, the fungus Marasmius oreades (known as the fairy ring champignon; see below) forms such a thick, water-repellent mass of hyphae beneath the soil surface that it prevents water from penetrating the soil and therefore leads to a ring of dead turf, with a ring of darker, lusher grass behind this. M. oreades is known to produce hydrogen cyanide in laboratory culture, and HCN also has been detected in the killing zone of rings. But there is uncertainty about whether this contributes to death of the turf. In fact, the simplest way of overcoming the killing action of this fungus is to water the affected zone of turf with a dilute solution of a surfactant such as washing-up liquid.

Ring of dead grass turf caused by growth of the fungus Marasmius oreades. The ring is expanding outwards towards the left of the image. A zone of dark, lush grass growth is seen behind the killing zone.

The same fairy ring as above, but part of the turf has been lifted to show the mass of water-repellent fungal hyphae growing on the dead roots and leaf sheaths of the turf.

Fruitbodies of Marasmius oreades. They are brown-coloured, with shallow caps (when fully expanded) and widely-spaced gills. The caps are about 3 cm diameter.