Basidiomycota

Sarah C Watkinson, University of Oxford, Oxford, England

Basidiomycota comprise the most morphologically complex group of macrofungi. They include mushrooms and toadstools, and rust and smut parasites of plants. Basidiomycetes grow as networks of hyphae colonizing nutrient substrates. Reproduction is by airborne spores produced from basidiomes such as mushrooms. Features used in identification show frequent convergent evolution classification is changing as molecular phylogeny reveals true relationships. Basidiomycetes are essential in carbon cycling in temperate and boreal forests, as wood decomposers and ectomycorrhizal symbionts. They form underground resource-sharing networks (the wood-wide web) which support plant biodiversity in forest ecosystems. Some form symbioses with animals, notably ants, termites, beetles and wasps. Others form lichens with photosynthetic microbes. Economically, mushrooms are valued for food and for bioactive compounds exploited in traditional medicine. Wood decay Basidiomycetes can destroy construction timber, but are valuable in lignocellulose conversion, for composting and as potential producers of biofuels. Rusts and smuts cause major crop losses.
. Life Cycle . Genetics

Advanced article
Article Contents
. Introduction

. Classification and Phylogeny . Ecology . Economic Importance

Online posting date: 15th December 2008

Introduction
Basidiomycota constitute a Phylum of the Kingdom Fungi that includes the most conspicuous and well-known fungi. There are about 35 000 known species, including all those that produce mushrooms and toadstools, jelly fungi, parasitic species causing rust and smut diseases on plants and yeast-like forms including the emerging human pathogen Cryptococcus neoformans. New species are continually being discovered. Diagnostic morphological features of basidiomycete fungi include dikaryotic vegetative mycelium, in which parental haploid nuclei remain paired but separate during growth; the basidia, specialized spore-producing cells in which both nuclear fusion and meiosis occur; and some ultrastructural features of the cell. However, there is no unique set of characters shared by all Basidiomycetes. The evolutionary relationships of the species of Basidiomycota are being elucidated by molecular phylogeny. Many of the morphological characters commonly used for identication show convergent evolution, having evolved separately in dierent lineages. Consequently, classication is in a state of ux. Current authorities are the ninth edition of Ainsworths Dictionary of Fungi (Kirk et al., 2001), and the classication of Fungi resulting from the AFTOL (http://aftol.org/) and Deep Hypha (http://ocid.nacse.org/

ELS subject area: Microbiology How to cite: Watkinson, Sarah C (December 2008) Basidiomycota. In: Encyclopedia of Life Sciences (ELS). John Wiley & Sons, Ltd: Chichester. DOI: 10.1002/9780470015902.a0000347.pub2

research/deephyphae/) programmes (Hibbett et al., 2007; Swann and Hibbett, 2008). It is in the assignment of orders to higher-level taxa, and in the basal lineages of the phylogenetic tree, that major changes are being made. Historically, many species have been given more than one name. Synonyms are listed in the Index Fungorum, at http:// www.indexfungorum.org/Names/Names.asp. Basidiomycota play essential roles in carbon cycling and the mineral nutrient dynamics of terrestrial ecosystems (Cairney, 2005). Their mycelium predominates among the microbial ora of nutrient-limited forest soils in temperate and boreal latitudes. Mycelium extending from decomposing logs and ectomycorrhizal roots forms extensive networks that scavenge and transport nutrients in the forest oor the so-called wood-wide web. Symbioses with plants and animals are common among Basidiomycetes. About 6000 species are known as ectomycorrhizal mutualistic symbionts. Close symbiotic associations exist between some species of decomposer basidiomycetes, and ants or termites that cultivate them. Basidiomycetes also form lichen associations with unicellular photosynthetic organisms. Basidiomycota are exploited in horticultural mushroom production, as producers of bioactive compounds, to degrade phenolic compounds in bioremediation, for lignocellulose processing, and in recreational mushroom collecting. Economic losses due to Basidiomycetes include the ravages of crop pathogens, and the decay of timber.

Life Cycle
The life cycle of Basidiomycota diers from those of plants and animals by the absence of morphologically
1

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

dierentiated gametes. Mating is by fusion of haploid individuals. No diploid zygote is immediately formed instead, a special cell type, the dikaryon, is produced. In a typical life cycle of Agaricales (mushroom-forming fungi), Figure 1, meiosis in a specialized cell, the basidium, leads to the formation and release of abundant haploid, typically uninucleate, airborne spores. When these land on a suitable moist surface they germinate to produce a septate mycelium with haploid nuclei of a single genotype, a monokaryon. For sexual reproduction this must encounter and fuse with a second monokaryotic mycelium of dierent genotype, so as to produce a dikaryotic mycelium populated by haploid nuclei of both parental genotypes. On fusion of monokaryons, a remarkable event occurs: nuclei from each monokaryon undergo mitotic divisions and migrate reciprocally throughout the joint mycelium, assorting

themselves into pairs consisting of one of each parental type. Paired haploid nuclei (conjugate nuclei) remain associated in each hyphal compartment. The dikaryon is thus functionally diploid. The dikaryotic state is maintained throughout the life of the mycelium, which may endure for hundreds of years (Smith et al., 1992). Specialized development at the growing tip of each lament prevents separation of the nuclear pairs during hyphal tip growth. The process is as follows. Simultaneous (conjugate) division of both nuclei is co-ordinated with septum formation. At mitosis and septum formation, a backwardly directed side-branch develops (a hook cell), into which one of the mitotic spindles is directed, allowing parallel orientation of both spindles. The side branch anastomoses with the main lament behind the newly developed septum so that one daughter

Monokaryotic hyphae of opposite mating-type meet and share nuclei in a dikaryon Spores land and germinate on damp surfaces

Dikaryotic mycelium has conjugate pairs of nuclei in each compartment

Dolipore' septum with central hole Microscopic haploid spores of assorted mating-type, released into the air

Clamp connection Hyphal knot Dikaryotic mycelium foraging for nutrient sources Mycelium colonizing a food source dead wood

Gill tissue

Spores

Meiosis in basidium on gill mating-type genes segregate. Each basidium produces four meiospores following meiosis

Sporophore begins to form from hyphal knot

Gill

Sporophore composed of dikaryotic hyphae

Developing sporophore, fed by foraging mycelium

Dikaryotic mycelium, composed of microscopic hyphae growing in soil or wood or as ectomycorrhiza on tree roots
Figure 1 The life cycle of a mushroom forming fungus. ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

nucleus from each spindle is positioned on each side of the septum. The side branch is then closed by a further septum to restore the integrity of the compartment. The small looped side branches remain at each septum after the lament tip has grown on, and are misleadingly termed clamp connections (Figure 1). They are diagnostic of dikaryotic basidiomycete mycelium, although not all species of basidiomycetes produce them. See also: Mitosis

Development of Basidiomata
Typically, Basidiomata can only develop from dikaryotic mycelium. In Agaricomycotina, basidiospores are produced and released from multicellular, highly dierentiated basidiomata, such as mushrooms, puballs, bracket fungi and resupinate forms that grow as attened layers of tissue. Basidiospores develop on a specialized hymenial surface, with its area increased by surface folds, pores or gills. Spores are continuously liberated. One basidiome can release billions of spores a day. Basidiomata can be formed at any time during the life of a mycelium, triggered by environmental conditions. Mushroom primordia rst appear as knots of hyphae and dierentiate early into regions destined to form the distinctive tissues of cap, gills and stalk (Moore, 1996). As the fruiting body enlarges, cell dierentiation accompanied by tissuespecic gene expression is maintained by the action of diusible morphogens. Stalk elongation and gill expansion are driven by intracellular hydrostatic pressures. Mushrooms can push up tarmac surfaces by metabolically increasing osmotic pressure within the stalk, each hypha of which is corseted by bands of inextensible chitin microbrils running transversely in the cell wall. Hyphae dierentiate during basidiome development, and three types are recognizable: generative hyphae which continue to grow, thick-walled skeletal hyphae which confer rigidity, and binding hyphae of limited growth that bind the tissue together. Basidiomata constructed of one, two or three hyphal types are termed mono-, di- and trimitic, respectively. The number of dierent hyphal types is used as a character for of the species identication, particularly in Polyporales (Ryvarden and Gilbertson, 1993/94). Metabolic changes during basidiome development include production of secondary metabolites the pigments, toxins and avours associated with mushrooms. There are hundreds of such molecules and many are of pharmacological interest (Ooi, 2001). See also: Mushrooms and Mushroom Cultivation The positioning of fruiting bodies is precisely regulated so that the spore-bearing surface is held vertical, allowing the spores to fall under gravity after discharge from the surface. Position is adjusted by tropic (bending) responses to gravity, and at some stages and species, to light direction.

body). The terminal cell of a hypha becomes delimited by a septum and enlarges into a club-shaped swelling. The two haploid nuclei of the dikaryon then fuse, and within minutes the diploid nucleus undergoes meiosis, producing a tetrad of daughter nuclei. Spurs sterigmata then develop at the end of the basidium, and one of each of the four nuclei resulting from meiosis moves through each sterigma, and becomes the nucleus of the spore which develops at its tip. Variations on this typical pattern occur, and some species have binucleate basidiospores. After spore formation, the basidium remains and contains a large vacuole. The basidium actively discharges all four basidiospores, by a mechanism involving the release of a uid drop at the point of attachment (Pringle et al., 2005). The energy released as this coalesces with the wet spore surface propels the spores just far enough to escape from the layer of basidia on the undersurface of the mushroom, and they then fall under gravity into the air beneath the fruiting body. Actively discharged ballistospores are widespread in Basidiomycetes. Not all basidiomycetes actively discharge spores, however. The polyphyletic group Gasteromycetes (including, e.g. puballs, earthstars and earthballs) produce basidiospores as a powder within a closed fruiting body, expelled out of holes or breaks in the outer skin when the puball is struck by raindrops. Basidia of Agaricomycetes are typically aseptate with short determinate sterigmata. Other groups have variable types of basidiospore formation and release. The basidia of jelly fungi such as Auriculariales and Tremellales are deeply septate, with long sterigmata and basidiospores which may bud to produce further spores. In temperate woodland in autumn, when most basidiomycetes sporophores appear, basidiospores may predominate in the air spora. Most spores are deposited close to the parent sporophore, but a few may travel for miles. See also: Fungal Spores Anamorphic spores are those formed asexually. The anamorphic states of basidiomycetes include the oidia formed on monokaryotic mycelium, yeast-like fungi including many that form ballistospores, and asexual phases of rust and smut life cycles.

Mating systems
Mushroom formation normally occurs only on dikaryotic mycelium produced by fusion between compatible monokaryons of dierent mating-type. Mating-type is determined by multiallelic factors at specic genetic loci, A and B, with compatibility between partners carrying dierent allelic forms. Typically, there are hundreds of dierent compatible mating-types in a population. Species with only one mating-type locus are termed bipolar, those with two are tetrapolar. Bipolarity appears to have evolved from tetrapolarity in several dierent lineages. Some species are capable of producing sporophores without mating.
3

Basidium structure and spore release

In Agaricomycetes, basidia develop on the hymenial surface of a basidiome (also termed a sporophore or fruiting

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

Coprinopsis cinereus illustrates the function of the two mating-type factors in a tetrapolar system (Casselton and Ku es, 2007). Each controls a dierent stage of dikaryon development. Heterozygosity at A allows the mycelium to develop hook cells for conjugate nuclear division. The A locus is a complex that encodes members of the homeobox family of transcription factors, which heterodimerize on mating to produce an active transcription factor. Heterozygosity at B allows regulated nuclear migration. The B locus encodes peptide pheromones and their receptors that are believed to mediate intracellular signalling between the two haploid nuclei of the dikaryotic hypha. Thus the A and B factors act together to maintain the dikaryotic state as the hypha grows by tip extension, septation and nuclear mitosis (Brown and Casselton, 2001). The tetrapolar multiallelic mating-type system maximizes outbreeding while reducing the possibility of sib mating. In the basidium, meiosis of a diploid nucleus of genotype A1A2B1B2 generates four haploid nuclei A1B1, A1B2, A2B1 and A2B2, each of which passes into one basidiospore. The chances of compatibility with another haploid from the same mushroom are thus 1 in 4. However, haploids from dierent basidiomata are almost certain to be compatible because within the population there are around a hundred dierent alleles at each of the A and B loci.

Mycelium
Although a basidiomycete may be identied by its conspicuous sporophores, it is the mycelium that persists and assimilates, mates and continually interacts with the environment, and thus corresponds to the biological concept of the individual organism. Mycelium is formed by apical extension, septation, branching, anastomosis and dierentiation of indeterminately extending hyphae. The mycelium of some basidiomycete individuals may reach metres in extent when growing in a low-disturbance environment. Single clones of Armillaria have been found throughout woodlands of many hectares (Smith et al., 1992). Many species that exploit habitats in which carbon sources and mineral nutrients are spatially separate develop cords (Boddy, 1999; Figure 2d) and rhizomorphs that channel long-distance transport of resources including water, air and carbon and mineral nutrients in solution, by mass ow. Mycelia in woodland, based on carbon-rich resources such as symbiotic roots or dead wood, can thus extend by hyphal tip growth to scavenge phosphate and nitrate from surrounding soil. Nutrient ow (translocation) occurs in cords at speeds of about 25 cm h21 and is responsive to source/sink gradients within the mycelium. Mathematical analysis of networks of cord forming wood decay fungi indicates that their architecture is adapted to function as an ecient resource supply

(a)

(b)

(c) 5 cm

3 cm (d) (e) (f)

Figure 2 (a) Wood decayed by a brown rot fungus leaving brown lignin-rich residue. (b) Wood partly decayed by a white rot fungus that has removed lignin but has not yet completely degraded the fibrous cellulose component of the wood. (c) A resupinate basidiome: the recently discovered Ceriporiopsis herbicola. Fortey & Ryvarden (Synopsis Fungorum (Oslo), 2007, vol. 23, p. 14, http://www.indexfungorum.org/, reproduced by kind permission of the Fungus Survey of Oxfordshire). (d) Mycelial cords and mycelium of a cord-forming wood decay fungus feeding on a fallen log in beech woodland (reproduced from Watkinson et al., 2006). (e) and (f) The ectomycorrhizal species Laccaria amethystea: (e) Basidiome; (f) Beech tree rootlet ensheathed in mycelium of the same species, identified by matching its 18S rRNA ITS sequence to that of the basidiome (Gardes and Bruns, 1993).

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

network (Fricker et al., 2007). For a given biomass, they appear to maximize conductivity, and robustness against loss of links caused by fungus-eating soil fauna (Boddy and Jones, 2007). Hyphae and mycelium provide fewer morphological clues to species than do sporophores. The ultrastructure of basidiomycete hyphae proves more consistent with phylogenetic relationships than their gross morphology. Phylogenetically consistent cellular structures include the type of septum, with its septal pore and associated parenthosomes, together termed the dolipore, and also the spindle pole body, an electron-dense structure that appears during nuclear division. The chemical composition of the hyphal wall polysaccharides also varies consistently in clades, with xylose present in all but Ustilaginomycetes (Hibbett and Thorn, 2001).

Classification and Phylogeny

Basidiomycota are a Phylum of the Subkingdom Dikarya of the Kingdom Fungi, and are classied in three Subphyla: Kingdom Fungi Subkingdom Dikarya Phylum Basidiomycota Subphylum Pucciniomycotina Ustilaginomycotina Agaricomycotina Only the commonest fungi in these groups are within the scope of this article; for a more detailed account see Hibbett et al. (2007) and Kirk et al. (2001). Classication of fungi is changing rapidly, as new understanding from molecular phylogeny is incorporated into schemes originally based on morphology. Current phylogenetic analysis of Basidiomycetes is based on multiple genes, mainly for ribosomal DNA, as well as mitochondrial and protein-coding genes tef1, rpb1, rpb2, atub and b-tub. Polyphyletic groups of previously unknown anity, particularly among yeast forms and jelly fungi, are thus being assigned according to their true evolutionary relationships. In Agaricomycotina, there has been extensive convergent evolution, particularly of types of basidiomata, not surprisingly as these are adapted solely for basidiospore production. Examples include basidiomes that are at (resupinate), as shown in Figure 2c (Binder et al., 2005), or have the shape of cups or tubes (cyphelloid) (Bodensteiner et al., 2004). It is unfortunate for mycologists that basidiomes are frequently the only structures available for eld identication. Because classication is currently being revised to accommodate new knowledge of the evolutionary relationships between genera, there is a problem reconciling new understanding of phylogeny with the traditional hierarchical categories of mycological taxonomy. Keys for eld identication must use available morphological characters. However, the techniques of molecular phyogeny have shown that many characters valuable for identication are not reliable for classication. See also: Evolution: Convergent and Parallel Evolution; Molecular Phylogeny Reconstruction

Genetics
In basidiomycetes, genotype determines not only mating compatibility as described earlier, but also the potential of mycelial individuals for vegetative fusion. Vegetative compatibility is determined by the vegetative compatibility (vc) loci of the fusing partners (Hansen and Hamelin, 1999). Unlike mating-type genes, identical alleles at all vc loci of both individuals are necessary for compatibility. Species vary in the number of vegetative compatibility groups (vcg) in the population. The larger the number of groups, the less likely are chance-met mycelia to be capable of fusion. Vegetative incompatibility is believed to limit the spread of selsh genetic elements by hyphal fusions between individuals in a population. See also: Intragenomic Conict Genetic methods enable basidiomycete fungi to be identied from environmental samples such as soil, air or water. This has enormously increased understanding of their population structure, ecosystem functions, and the origin of pests and pathogens (e.g. Taylor and Fisher, 2003). Genetic strain markers include polymorphic forms of matingtype and vegetative compatibility genes, as well as the more variable parts of the genome such as microsatellite deoxyribonucleic acid (DNA). The internal transcribed spacer (ITS) region of nuclear rDNA is widely used for species identication (Gardes and Bruns, 1993). See also: Molecular Ecology Population genetics can elucidate the geographical origin of strains and species. Analysis of the worldwide distribution of strains of the building dry rot fungus Serpula lacrymans, based on polymorphisms in mating-type and vegetative compatibility loci, showed that it has evolved relatively recently from a wild relative, probably in Asia, and subsequently spread rapidly within the built environment (Kauserud et al., 2007). Basidiomycete genetics is beneting from the publication of whole-genome sequences of a number of species including plant parasites, ectomycorrhizal fungi and wood decomposers (e.g. Martinez et al., 2004; Martin et al., 2008).

Pucciniomycotina
The most numerous group is the order Pucciniales (formerly Uredinales or the rust fungi) with about 7000 species in 14 families and 163 genera (Kirk et al., 2001). They are highly host-specic obligate biotrophic parasites on leaves and stems of land plants, including ferns, gymnosperms and angiosperms (Frieders et al., 2008). Their dikaryotic mycelium grows intercellularly, usually with haustoria. Up to ve spore stages are produced. Throughout summer, rusty red dikaryotic urediniospores spread rust disease, causing epidemics in crops. Teliospores produced in late summer are survival spores, from which a short mycelium
5

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

develops on germination, producing the septate 24-celled basidia. Many rust species are heteroecious, alternating between two dierent plant host species.

Heterobasidiomycetes
These include the jelly fungi, a disparate group with unusual basidiomata, often with simple nondolipore septa, and septate basidia with indeterminately growing sterigmata. Genera with conspicuous basidiomata familiar to eld mycologists include Tremella, with gelatinous basidiomata, typically parasitic on other wood decomposing fungi, Dacrymyces, strongly cellulolytic decomposers of wet wood, and Auricularia, for example Auricularia auricula-judae saprobic on Sambucus niger. The phylogenetic relationships of these fungi within Basidiomycota are being gradually elucidated.

Ustilaginomycotina
The most numerous group is the order Ustilaginales (smut fungi) with 672 species in 2 families and 14 genera. They parasitize plants, mainly Gramineae and Cyperaceae, and are host-specic obligate biotrophs in the parasitic phase (Bauer et al., 2008). However they grow as saprotrophicbudding unicells in culture. Their diploid, uninucleate spores typically form a dark powder conspicuous on or beneath the surface of host leaves or owers. Meiosis occurs on germination, with the production of a septate basidium bearing four basidiospores.

Ecology
The most important roles of basidiomycetes in ecosystems are as wood decomposers, as symbionts of plants and insects, and as plant parasites. See also: Fungal Ecology

Agaricomycotina
The Agaricomycotina comprise three orders, Tremellomycetes, Dacrymycetes and the Agaricomycetes which include the majority of known species (Hibbett et al., 2007). Agaricales, the gilled fungi (mushrooms and toadstools) include 9387 described species. Well-known examples are Coprinus (ink caps) and Agaricus (including the cultivated mushrooms Agaricus bisporus), as well as the cultivated edible species Lentinus edodes (shi-itake) and the oyster mushroom Pleurotus ostreatus. The genus Amanita includes the deadly death cap, Amanita phalloides, which produces several animal toxins including amanitin, which inhibits the biosynthesis of messenger ribonucleic acid (RNA). Armillaria, a tree parasite that produces extensive systems of rhizomorphs in woodland, includes Armillaria bulbosa, reported as the largest and oldest living organism (Smith et al., 1992). See also: Fungal Metabolites; Mushrooms and Mushroom Cultivation The Polyporales have 2253 species and include many wood decomposing fungi with bracket-shaped basidiomata (Ryvarden and Gilbertson, 1993/94). The genus Ganoderma includes the medicinal fungus Ganoderma lucida (reishi), and the spectacular Ganoderma applanatum, Dryads saddle with brackets up to half a metre in diameter.

Wood decomposing fungi

Basidiomycete are the main wood decomposers of fallen woody litter in temperate and boreal zones, and thus play an essential role in the carbon cycle on land. Most species degrade both lignin and cellulose, causing white rot (Figure 2b). The enzymes involved in lignin degradation include laccases, manganese peroxidases and lignin peroxidases (Harvey and Thurston, 2001). Lignin peroxidases acting on hydrogen peroxide produced in metabolism generate highly reactive hydroxyl free radicals. These react with the complex aromatic lignin molecule to produce a short-lived unstable aromatic free radical which breaks into a variety of molecular fragments. The process is oxidative and has been described as enzymatic combustion. The whole genome of the lignin-degrading wood decay fungus Phanerochaete chrysosporium has recently been sequenced (Martinez et al., 2004), revealing many genes encoding secreted oxidases, peroxidases and hydrolytic enzymes that cooperate in wood decay. Lignin is not itself utilizable as a carbon or energy source by the fungus but its removal unmasks utilizable cellulose. Brown rot wood decay, Figure 2a, caused by relatively few species, and occurring mainly on conifer wood, is the commonest type of decay in boreal coniferous forests, generating lignin-rich residues which comprise up to 30% of soil sequestered carbon (Ryvarden and Gilbertson, 1993/ 94). Cord-forming wood decay fungi (Boddy, 1999) form extensive forest oor networks which scavenge and redistribute mineral nutrients (Watkinson et al., 2006). See also: Global Carbon Cycle; Lignins; Soils and Decomposition; Xylem Structure and Function

Basidiomycete yeasts
There are about 220 species of yeasts that have basidiomycete characteristics such as the production of ballistospores. They occur in each of the three subphyla of Basidiomycota (Fell et al., 2001). Sporobolomyces includes common asexual yeasts which discharge ballistospores that multiply by budding and are widespread on the surface of leaves and fruits. The human pathogen Cryptococcus neoformans is the asexual phase of Filobasidiella neoformans, causing opportunistic cryptococcosis, mainly in immunocompromized individuals, who become infected by inhaling spores. See also: Fungal Pathogens of Humans; Mycoses
6

Mutualistic symbioses
Plant symbioses
Ectomycorrhiza are symbiotic associations formed between basidiomycete mycelium and the roots of living

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

trees (Smith and Read, 1997). Root and mycelium become closely intermingled, with a fungal sheath enclosing the root tip and hyphae penetrating between the cortical cells of the root. The ectomycorrhizal morphotype is characteristic of each association of species (Cairney and Chambers, 1999; Figure 2e and f). The fungal mycelium acquires carbon compounds for growth from the host tree, and the tree acquires plant mineral nutrients, particularly phosphate, scavenged from soil and litter by fungal mycelium (Read and Perez-Moreno, 2003). Ectomycorrhizal connections can channel carbon compounds between trees of dierent species, and can supply photosynthate to seedlings in which photosynthesis is reduced by shading (Simard et al., 1997). These results suggest a prominent role for mycorrhizal basidiomycete mycelium in resource allocation within plant communities (Lindahl et al., 2002), encapsulated by the term wood-wide web (Read, 1997). Tree species in Betulaceae, Dipterocarpaceae, Fagaceae, Pinaceae and Myrtaceae form ectomycorrhizal associations. Resource sharing between plants connected by ectomycorrhizal mycelium can enable new plant species to become established on bare soil (Nara and Hogetsu, 2004). Ectomycorrhizal symbiosis is evolutionarily unstable, having been gained and lost in various basidiomycete lineages (Hibbett et al., 2000). Some plants that lack chlorophyll grow as highly host-specic parasites on the underground mycelium of basidiomycetes (Bidartondo and Bruns, 2002). See also: Mycorrhiza

regions of the world. There are around 7000 rust species and many of these aect crop plants and have worldwide distribution. Control is mainly by breeding for resistance, both by incorporating specic rust resistance genes into cultivars, and also selecting for multigenic resistance. With normal control measures (breeding resistant cultivars, crop hygiene) the corn smut Ustilago maydis causes annual losses of 2% on average, but up to 100% can be lost in local, badly aected areas. As well as the rusts and smuts that infect above-ground parts of plants, the microscopic rootinfecting Rhizoctonia is also a basidiomycete. It is widespread in soil, and attacks seedlings and roots. The wood decaying basidiomycete Heterobasidion annosum causes losses in forestry plantations. Fungi in the genus Armillaria are serious tree pathogens that spread in plantations on sites where felling of preexisting infected woodland has left residual mycelium in stumps. It has broad host specicity and its rhizomorphs invade the roots of adjacent healthy trees. See also: Fungal Pathogens of Plants

Timber decay
Wood decaying basidiomycetes are the principal cause of decay of construction timber, in buildings, railway sleepers and transmission poles. Dry rot in buildings is caused by the brown rot fungus Serpula lacrymans which can spread aggressively within a building, by translocating water and nutrients rapidly through mycelial cords across masonry from its foodbase in decaying timber to supply the advancing mycelium (Money, 2006). Other genera that commonly decay building timbers include species of Coniophora, Trametes, Lentinus and Gloeophyllum. Because fungal decay spores are ubiquitous in air, any damp timber is liable to decay, so timber preservation is of major industrial importance. The lignin-degrading enzymes of white rot basidiomycetes are exploited in bioremediation to break down pollutant aromatic compounds.

Animal symbioses
Basidiomycetes have many relationships with insects. Mushroom fruiting bodies may be home to hundreds of species of dipteran larvae, and those of bracket fungi can support hundreds of species of beetle larvae, with some specic to the fungal species. Septobasidiales are a group which associate with scale insects on leaves in a tripartite relationship. Attine ants and termites have both evolved symbiosis with basidiomycete species in the order Lepiotales (Mueller et al., 2005). The workers feed the fungal mycelium with vegetation gathered from the neighbourhood of the nest, and the migrating queen carries the fungal culture to new nests. Molecular analysis has shown that in one case, a single fungal clone has been maintained for 23 million years by a single ant species. Symbiosis with the fungus enables the ants to feed on a wide variety of plant material which is converted into nutrient-rich fungal food by the mycelium. See also: Mutualistic Symbioses

Mushroom production
Hundreds of thousands of tonnes of the cultivated mushroom, Agaricus bisporus, are produced annually in Europe, and other species including the oyster mushroom Pleurotus ostreatus and shi-itake, Lentinus edodes, are also important crops. Many species of wild mushrooms are also commercially available, some such as matsutake, the Asian ectomycorrhizal species Tricholoma matsutake being particularly prized (Moore and Chiu, 2001). See also: Mushrooms and Mushroom Cultivation

Economic Importance
Crop and forest pathogens
The wheat stem rust fungus Puccinia graminis causes losses in North America of about 1 million tonnes of wheat annually, up to 100 million in years of severe epidemics, and these gures are typical of those in other wheat growing

References
Bauer R, Begerow D and Oberwinkler F (2008) Ustilaginomycotina, Ustilaginomycetes, the true smut fungi. Bidartondo MI and Bruns TD (2002) Fine-level mycorrhizal specicity in the Monotropoideae (Ericaceae): specicity for fungal species groups. Molecular Ecology 11: 557569.

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

Binder M, Hibbett DS, Larsson K-E et al. (2005) The phylogenetic distribution of resupinate forms across the major clades of mushroom-forming fungi (Homobasidiomycetes). Systematics and Biodiversity 3: 113157. Boddy L (1999) Saprotrophic cord forming fungi: meeting the challenge of heterogeneous environments. Mycologia 91: 1332. Boddy L and Jones TH (2007) Interactions between Basidiomycota and invertebrates. In: Boddy L, Frankland JC and van west P (eds) Ecology of Saprotrophic Basidiomycetes, pp. 155179. London: Academic Press. Bodensteiner PM, Binder M, Moncalvo J-M, Agerer R and Hibbett DS (2004) Phylogenetic relationships of cyphelloid homobasidiomycetes. Molecular Phylogenetics and Evolution 33: 501515. Brown AJ and Casselton LA (2001) Mating in mushrooms: increasing the chances but prolonging the aair. Trends in Genetics 17: 393400. Cairney JWG (2005) Basidiomycete mycelia in forest soils: dimensions, dynamics and roles in nutrient distribution. Mycological Research 109: 720. Cairney JWG and Chambers SM (eds) (1999) Ectomycorrhizal Fungi: Key Genera in Prole. Berlin: Springer. Casselton LA and Ku es U (2007) The origins of multiple mating types in the model mushrooms Coprinopsis cinerea and Schizophyllum commune. In: Heitman J, Kronstad JW, Taylor JW and Casselton LA (eds) Sex in Fungi: Molecular Determination and Evolutionary Implications. Washington, DC: ASM Press. Fell JW, Boekhout T, Fonseca A and Sampaio JP (2001) Basidiomycetous yeasts. In: McLaughlin DJ, McLaughlin EJ and Lemke P (eds) The Mycota. Vol. VII. Part B., Systematics and Evolution, pp. 335. Berlin: Springer. Fricker MD, Boddy L and Bebber DP (2007) Network organisation of mycelial fungi. In: Howard RJ and Gow NAR (eds) The Mycota, vol. 8, Biology of the fungal cell, 2nd edn, pp. 309 330. Berlin: Springer-Verlag. Frieders EM, McLaughlin DJ and Szabo LJ (2008) Pucciniomycotina, Urediniomycotina. A diverse group including rusts, yeasts, smut-like and jelly like fungi. http://tolweb.org/ Pucciniomycotina/20528 Gardes M and Bruns TD (1993) ITS primers with enhanced specicity for basidiomycetes: application to the identication of mycorrhizae and rusts. Molecular ecology 2: 113118. Hansen EM and Hamelin RC (1999) Population structure of basidiomycetes. In: Worrall JJ (ed.) Structure and Dynamics of Fungal Populations, chap. 11, pp. 251281. Dordrecht: Kluwer Academic Publishers. Harvey PJ and Thurston CF (2001) The biochemistry of ligninolytic fungi. In: Gadd GM (ed.) Fungi in Bioremediation, chap. 2, pp. 2751. Cambridge: Cambridge University Press. Hibbett DS and Thorn RG (2001) Homobasidiomycetes. In: McLaughlin DJ, McLaughlin EJ and Lemke P (eds) The Mycota. Vol. VII. Part B., Systematics and Evolution, pp. 121168. Berlin: Springer. Hibbett DS, Gilbert L-B and Donoghue MJ (2000) Evolutionary instability of ectomycorrhizal symbioses in basidiomycetes. Nature 407: 506508. Hibbett DS, Binder M, Bischo JF et al. (2007) A higher-level phylogenetic classication of the Fungi. Mycological Research 111: 509547.

rden IB, Stre G-P et al. (2007) Asian origin Kauserud H, Svega and rapid global spread of the destructive dry rot fungus Serpula lacrymans. Molecular Ecology 16: 33503360. Kirk PM, Cannon PF, David JC and Stalpers JA (eds) (2001) Ainsworth and Bisbys Dictionary of the Fungi, 9th edn, 655 pp. Wallingford, UK: CAB International. Lindahl BO, Taylor AFS and Finlay RD (2002) Dening nutritional constraints on carbon cycling in boreal forests towards a less phytocentric perspective. Plant and Soil 242: 123135. n D et al. (2008) The genome of Laccaria Martin F, Aerts A, Ahre bicolor provides insights into mycorrhizal symbiosis. Nature 452: 8892. Martinez D, Larrondo LF, Putnam N et al. (2004) Genome sequence of the lignocellulose degrading fungus Phanerochaete chrysosporium strain RP78. Nature Biotechnology 22: 695700. Money NP (2006) Plagues upon houses and cars: the unnatural history of Meruliporia incrassata, Serpula lacrymans and Spaherobolus stellatus. In: Gadd GM, Watkinson SC and Dyer PS (eds) Fungi in the Environment, pp. 289309. Cambridge: Cambridge University Press. Moore D (1996) Inside the developing mushroom cells, tissues and tissue patterns. In: Chiu S-W and Moore D (eds) Patterns in Fungal Development, chap. 1, pp. 136. Cambridge: Cambridge University Press. Moore D and Chiu SW (2001) Fungal products as food. In: Pointing SB and Hyde KD (eds) Bio-Exploitation of Filamentous Fungi, chap. 10, pp. 223251. Hong Kong: Fungal Diversity Press. Mueller UG, Gerardo NM, Aanen DK, Six DL and Schulz T (2005) The evolution of agriculture in insects. Annual Review of Ecology, Evolution and Systematics 36: 563595. Nara K and Hogetsu T (2004) Ectomycorrhizal fungi on established shrubs facilitate subsequent seedling establishment of successional plant species. Ecology 85: 17001707. Ooi VEC (2001) Higher fungi in traditional Chinese medicine. In: Pointing SB and Hyde KD (eds) Bio-Exploitation of Filamentous Fungi, chap. 9, pp. 191222. Hong Kong: Fungal Diversity Press. Pringle A, Patek SN, Fischer M, Stolze J and Money NP (2005) The captured launch of a ballistospore. Mycologia 97: 866871. Read DJ (1997) Mycorrhizal fungi: the ties that bind. Nature 388: 517518. Read DJ and Perez-Moreno J (2003) Mycorrhizas and nutrient cycling in ecosystems a journey towards relevance? New Phytologist 157: 475492. Ryvarden L and Gilbertson RL (1993/94) European Polypores, vol. 1 (1993), vol. 2 (1994) Oslo: Fungiora. Simard SW, Perry DA, Jones MD et al. (1997) Net transfer of carbon between ectomycorrhizal tree species in the eld. Nature 388: 579582. Smith ML, Bruhn JN and Anderson JB (1992) The fungus Armillaria bulbosa is among the largest and oldest living organisms. Nature 356(2): 428431. Smith SE and Read DJ (1997) Mycorrhizal Symbiosis. London: Academic Press. Swann E and Hibbett DS (2008) Basidiomycota. The club fungi. http://tolweb.org/Basidiomycota/20520 Taylor JW and Fisher MC (2003) Fungal multilocus sequence typing its not just for bacteria. Current Opinion in Biology 6: 351356.

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net

Basidiomycota

Watkinson SC, Bebber D, Darrah PR et al. (2006) The role of wood decay fungi in the carbon and nitrogen dynamics of the forest oor. In: Gadd GM (ed.) Fungi in Biogeochemical Cycles. Cambridge: Cambridge University Press.

Further Reading
Boddy L, Frankland JC and van West P (2007) Ecology of Saprotrophic Basidiomycetes. Amsterdam: Academic Press/ Elsevier. Breitenbach J and Kranzlin F. Fungi of Switzerland vols 25 (19862000); Kranzlin F Fungi of Switzerland vol. 6 (2005). Lucerne: Verlag Mykologia.

Cannon PF and Kirk PM (2007) Fungal Families of the World. Wallingford: CABI Publishing. Carlile MJ, Gooday GW and Watkinson SC (2001) The Fungi, 2nd edn. London: Elsevier. Grin DH (1994) Fungal Physiology, 2nd edn. New York: WileyLiss. Phillips R (2006) Mushrooms. London: Macmillan. Rayner ADM and Boddy L (1988) Fungal Decomposition of Wood. Chichester, UK: Wiley. Spooner B (2005) Collins Wild Guide: Mushrooms and Toadstools. London: Harper Collins. Webster J (1980) Introduction to Fungi, 2nd edn. Cambridge: Cambridge University Press.

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net