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Basidiomycota (Club Fungi)  the Basidiomycota (Colloquially Basidiomycetes) Are a Large Group of Fungi with Over 30 000 Species

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Basidiomycota (Club Fungi)  the Basidiomycota (Colloquially Basidiomycetes) Are a Large Group of Fungi with Over 30 000 Species B.Sc Botany (Hons) Semester II Mycology and Phytopathology Basidiomycota (club fungi) The Basidiomycota (colloquially basidiomycetes) are a large group of fungi with over 30 000 species. They include many familiar mushrooms and toadstools, bracket fungi, puffballs, earth balls, earth stars, stinkhorns, false truffles, jelly fungi and some less familiar forms. Also classified here are the rust and smut fungi, which are pathogens of higher plants and may cause serious crop diseases. Most basidiomycetes are terrestrial with wind-dispersed spores, but some grow in freshwater or marine habitats. Many are saprotrophic and are involved in litter and wood decay, but there are also pathogens of trees. the honey fungus, Armillaria, which attacks numerous tree species, and Heterobasidion annosum, which can seriously damage conifer plantations. 1. The somatic phase consists of a well developed, septate, filamentous mycelium which passes chiefly through two stages: a) primary mycelium- it is formed by the germination of a basidiospore and contains a single haploid nucleus in each cell. It bears neither sex organs nor any basidia and basidiospores. It is short lived b) Secondary or dikaryotic mycelium It constitutes the main food absorbing phase and consists of cells each containing two haploid nuclie. It is long lived and plays prominent role in the life cycle. In Homobasidiomycetidae it may continue to grow for years producing fructifications every year by the interweaving of hyphae. The fructifications bear basidia and basidiospores In Heterobasidiomycetidae it forms teleutospores or brand spores which germinate to produce basidia bearing basidiospores. 2.Except in rusts and smuts the septal pore in the Basidiomycetes is complex, It is dolipore septa with parenthesome type. 3.The motile cells are absent in the life cycle 4. The clamp connections on the dikaryotic hyphae are of universal occurrence 5.Asexual reproduction by spores plays an insignificant role in the life cycle. The homobasidiomycetes do not form any asexual spores. The heterobasidiomycetes form them in the dikaryotic mycelium and produces uredospores and aecidiospores in the rust. 6.The sex organs are lacking in the Basidiomycetes. The sexual process is represented by plasmogamy and karyogamy. Karyogamy is immediately followed by meiosis. 7. Basidium is the characteristic reproductive organ of Basidiomycetes in which both karyogamy and meiosis takes place. 8.Typically the basidium bears four basidiospores exogenously. The number, however varies from one to many depending on species. 9. The basidiospore germinates to produce the primary mycelium. Resembles with Ascomycota 1. Both include parasitic as well as saprophytic species. 2. Terrestrial mycelium consists of septa. 3. The septa have each a central pore. 4. Motile cells completely absent. 5. The sexual process comprises plasmogamy and karyogamy. The latter is immediately followed by meiosis. 6. The delayed fusion of nuclei of opposite strains after plasmogamy has resulted in the origin and establishment of a binucleate or dikaryophase in the life cycle. 7. The characteristic reproductive organ, basidium of Basidiomycetes and ascus of Ascomycetes resemble each other in development and cytology till the initiation of spores. Both arises from binucleate cells, the basidium from the dikaryotic hyphae and ascus from the ascogenous hyphae. Karyogamy and meiosis both occur in the basidium as in the ascus. 8.The basidiospores and ascospores are ususally haploid and uninucleate. 9. The basidiocarp in Basidiomycetes and ascocarp in Ascomycetes (not homologous structure) 10.A clamp connection of the Basidiomycetes is considered homologous in sturcture and analogous in function to the hook of the Ascomycetes. The Basidiomycetes differ from the Ascomycetes in the following respects: 1. The septal pore in most of the Basidiomycetes is not a simple pore as in Ascomycetes but is a complex structure known as dolipore. The actual pore is barrel shaped. It is surrounded by a swollen rim which is a part of the annular septum. The opening of the pore is guarded by a curved pore cap called parenthesome. 2. The primary mycelium is short lived whereas in Ascomycetes it is long lived and dominanat. 3. The primary mycelium in the Basidiomycetes bears neither the sex organs nor basidia or basidiospores. 4. The conidia (uredospores and aeciospores) are borne on the secondary mycelium whereas in Ascomycetes they are borne on haplomycelium 5. Except the Uredinales all traces of sexual apparatus have been lost throughout the classes. 6. Presence of clamp connection is a characteristic feature of secondary mycelium 7. The dikaryotic (secondary) mycelium is long lived, independent structure whereas ascogenous hypae are short lived, homologous and occur inside the fruit body. 8.The fruit bodies in Basidiomycetes consists entirely of dikaryotic hyphae whereas in Ascomycetes the basal hyphae, peridia and paraphyses are haploid. 9.The basidium produced 4 basidiospores whereas ascus produced 8 ascospores. 10. The basidiospores are produced exogenously whereas ascospores are produced endogenously. 1. Homobasidiomycetes. Fungi with holobasidia, e.g. Agaricus and Polypores. 2.Heterobasidiomycetes. Fungi with heterobasidia, i.e. jelly fungi and their allies. 3. Urediniomycetes. Rust fungi. 4. Ustilaginomycetes. Smut fungi. Mycelium of Basidiomycetes • Heterothallic Basidiomycetes has 3 types: 1. Primary mycelium 2. Secondary mycelium 3. Tertiary mycelium 1. Primary mycelium: arises from germination of a single haploid basidiospores 1st mycelium to form At germination repeated mitotic nuclear division occurs. Early germ tubes become multinucleate and coenocytic. Transverse septa are laid down behind the growing hyphal tip and eventually divide the hypha into segments that are uninucleate. Uninucleate hypha are called monokaryotic . Such mycelia are said to be homokaryotic (Gr. homos=equal, alike; karyon= nut, here meaning nucleus) as they contain only one type of nucleus. This kind of mycelium in most Basidiomycetes is capable of indefinite growth but characteristically it develops into secondary mycelium. 2. Secondary Mycelium Forms when two basidiospores germinate near each other and the primary monokaryotic, mycelia of these genetically distinct mating types anastomose. fuse together and initiate the formation of a mycelium made up of segments, each containing two genetically distinct nuclei. Such mycelia are said to be dikaryotic and heterokaryotic. The dikaryotic mycelium will continue to grow, but the new growth will differ. 3. Tertiary mycelium arises when secondary mycelium transforms into organized, specialized tissues that comprise the basidiocarps of the most complex species. The hyphae constituting the basidiocarp may be differentiated several types. • Dikaryotization (diploidisation in older literature) conversion of monokaryon to a dikaryon Methods of dikaryotisation a) if the two monokaryotic strains are fully compatible, the mycelia make contact. The hyphal walls separating them break down and cytoplasmic continuity is established between them. Nuclei from one strain migrate rapidly establishing a heterokaryotic , binucleate cell. The binucleate cell produces a branch into which the nuclear pair migrates The two nuclei divide conjugately repeatedly and the sister nuclei separate as extensive septate mycelium. Migration is accompanied by nuclear division, in order to permit the doubling of the number of nuclei in both mycelia b). In another very common method of dikaryotisation, there a division of the nuclei in the above formed binucleate cell followed by migration of the daughter nuclei into primary mycelium of the opposite mating type, eg. The ‘a’ nucleus moves into ‘b’ mycelium, while a ‘b’ nucleus moves into the ‘a’ mycelium. The foreign nucleus in each mycelium then divides rapidly and its progeny migrating from one cell to another completing dikaryotising both parent mycelia c) Dikaryotisation can occur by the deposition of an oidium from one strain close to the mycelium of another. A hypha from the mycelium shows chemotropic curvature towards the oidium and fuses with it If the strains are compatible, nuclear migration follows, which associated with the breakdown of dolipore/parenthesome complex leading to the transfer of a compatible nucleus from one compartment to another. Eg. Coprinus cinereus d) Dikaryotisation can also occur if a monokaryon encounters a dikaryon carrying nuclei of a compatible mating type. Migration is unilateral instead of reciprocal Eg. Autoecious rusts Clamps connection In most Basidiomycetes, the most conspicuous feature of the secondary mycelium is the presence of clamp connections. Visible as a lateral bulge in the hyphal wall adjacent to a transverse septum. Clamp connections are special devices for maintaining the dikaryotic condition as growth continues. Ensured each segment of a dikaryotic hypha contains two genetically distinct nuclei. In the terminal segment of a dikaryotic hypha, a clamp connection develops between the position of a pair of nuclei ‘a’ and ‘b’ following the sequence The apical cell produces a short backward pointing side branch or a hook (clamp) and a daughter nucleus moves into it, the other nucleus remaining in the hypha but close to the branch. Simultaneous nuclear division follows One division becomes oriented obliquely so that one daughter nucleus a forms in the clamp connection and the other a’ in the dividing cell Division
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