89-468: Neolecta is a genus of ascomycetous fungi. The species share the English designation " Earth tongues " along with some better-known fungi (e.g. Geoglossum , Microglossum ) with a similar general form, but in fact they are only distantly related. Neolecta is the only genus belonging to the family Neolectaceae , which is the only family belonging to the order Neolectales . Neolectales , in turn,
178-470: A bipolar ( unifactorial ) or a tetrapolar ( bifactorial ) mating system. This results in the fact that following meiosis , the resulting haploid basidiospores and resultant monokaryons, have nuclei that are compatible with 50% (if bipolar) or 25% (if tetrapolar) of their sister basidiospores (and their resultant monokaryons) because the mating genes must differ for them to be compatible. However, there are sometimes more than two possible alleles for
267-408: A dikaryon . The hyphae are then said to be dikaryotic. Conversely, the haploid mycelia are called monokaryons . Often, the dikaryotic mycelium is more vigorous than the individual monokaryotic mycelia, and proceeds to take over the substrate in which they are growing. The dikaryons can be long-lived, lasting years, decades, or centuries. The monokaryons are neither male nor female. They have either
356-1196: A thallus usually referred to as the mycelium , which—when visible to the naked eye (macroscopic)—is commonly called mold . During sexual reproduction, many Ascomycota typically produce large numbers of asci . The ascus is often contained in a multicellular, occasionally readily visible fruiting structure, the ascocarp (also called an ascoma ). Ascocarps come in a very large variety of shapes: cup-shaped, club-shaped, potato-like, spongy, seed-like, oozing and pimple-like, coral-like, nit-like, golf-ball-shaped, perforated tennis ball-like, cushion-shaped, plated and feathered in miniature ( Laboulbeniales ), microscopic classic Greek shield-shaped, stalked or sessile. They can appear solitary or clustered. Their texture can likewise be very variable, including fleshy, like charcoal (carbonaceous), leathery, rubbery, gelatinous, slimy, powdery, or cob-web-like. Ascocarps come in multiple colors such as red, orange, yellow, brown, black, or, more rarely, green or blue. Some ascomyceous fungi, such as Saccharomyces cerevisiae , grow as single-celled yeasts, which—during sexual reproduction—develop into an ascus, and do not form fruiting bodies. In lichenized species,
445-474: A basidium). In some smuts such as Mycosarcoma maydis the nuclei migrate into the promycelium that becomes septate (i.e., divided into cellular compartments separated by cell walls called septa ), and haploid yeast-like conidia/basidiospores sometimes called sporidia, bud off laterally from each cell. In various smuts, the yeast phase may proliferate, or they may fuse, or they may infect plant tissue and become hyphal. In other smuts, such as Tilletia caries ,
534-472: A bottle shaped cell called a phialide , from which the spores are produced. Not all of these asexual structures are a single hypha. In some groups, the conidiophores (the structures that bear the conidia) are aggregated to form a thick structure. E.g. In the order Moniliales, all of them are single hyphae with the exception of the aggregations, termed as coremia or synnema. These produce structures rather like corn-stokes, with many conidia being produced in
623-442: A dikaryon. The dikaryon is long lasting but ultimately gives rise to either fruitbodies with basidia or directly to basidia without fruitbodies. The paired dikaryon in the basidium fuse (i.e. karyogamy takes place). The diploid basidium begins the cycle again. Coprinopsis cinerea is a basidiomycete mushroom. It is particularly suited to the study of meiosis because meiosis progresses synchronously in about 10 million cells within
712-413: A dimorphic, half yeast , half filamentous genus parasitic on leaves, branches, and catkins ; Schizosaccharomyces , a genus of fission yeasts (e.g. Schizosaccharomyces pombe ); and Pneumocystis , a yeast-like genus of mammalian parasites. To date, the genus has been unculturable, suggesting it is either obligately parasitic or symbiotic. It provides important evidence for the evolutionary history of
801-506: A double-dividing wall with a central lamella (layer) forms between the cells; the central layer then breaks down thereby releasing the spores. In rhexolytic dehiscence, the cell wall that joins the spores on the outside degenerates and releases the conidia. Several Ascomycota species are not known to have a sexual cycle. Such asexual species may be able to undergo genetic recombination between individuals by processes involving heterokaryosis and parasexual events. Parasexuality refers to
890-433: A form of pneumonia . Asci of Ascosphaera fill honey bee larvae and pupae causing mummification with a chalk-like appearance, hence the name "chalkbrood". Yeasts for small colonies in vitro and in vivo , and excessive growth of Candida species in the mouth or vagina causes "thrush", a form of candidiasis . The cell walls of the ascomycetes almost always contain chitin and β-glucans , and divisions within
979-476: A further mitotic division that results in eight nuclei in each ascus. The nuclei along with some cytoplasma become enclosed within membranes and a cell wall to give rise to ascospores that are aligned inside the ascus like peas in a pod. Upon opening of the ascus, ascospores may be dispersed by the wind, while in some cases the spores are forcibly ejected form the ascus; certain species have evolved spore cannons, which can eject ascospores up to 30 cm. away. When
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#17327733168761068-436: A given locus, and in such species, depending on the specifics, over 90% of monokaryons could be compatible with each other. The maintenance of the dikaryotic status in dikaryons in many Basidiomycota is facilitated by the formation of clamp connections that physically appear to help coordinate and re-establish pairs of compatible nuclei following synchronous mitotic nuclear divisions. Variations are frequent and multiple. In
1157-743: A hypha. Vegetative hyphae of most ascomycetes contain only one nucleus per cell ( uninucleate hyphae), but multinucleate cells—especially in the apical regions of growing hyphae—can also be present. In common with other fungal phyla, the Ascomycota are heterotrophic organisms that require organic compounds as energy sources. These are obtained by feeding on a variety of organic substrates including dead matter, foodstuffs, or as symbionts in or on other living organisms. To obtain these nutrients from their surroundings, ascomycetous fungi secrete powerful digestive enzymes that break down organic substances into smaller molecules, which are then taken up into
1246-529: A large-scale specialized structure that helps to spread them. These two basic types can be further classified as follows: Sometimes the conidia are produced in structures visible to the naked eye, which help to distribute the spores. These structures are called "conidiomata" (singular: conidioma ), and may take the form of pycnidia (which are flask-shaped and arise in the fungal tissue) or acervuli (which are cushion-shaped and arise in host tissue). Dehiscence happens in two ways. In schizolytic dehiscence,
1335-433: A mass from the aggregated conidiophores. The diverse conidia and conidiophores sometimes develop in asexual sporocarps with different characteristics (e.g. acervulus, pycnidium, sporodochium). Some species of ascomycetes form their structures within plant tissue, either as parasite or saprophytes. These fungi have evolved more complex asexual sporing structures, probably influenced by the cultural conditions of plant tissue as
1424-552: A separate artificial phylum , the Deuteromycota (or "Fungi Imperfecti"). Where recent molecular analyses have identified close relationships with ascus-bearing taxa, anamorphic species have been grouped into the Ascomycota, despite the absence of the defining ascus. Sexual and asexual isolates of the same species commonly carry different binomial species names, as, for example, Aspergillus nidulans and Emericella nidulans , for asexual and sexual isolates, respectively, of
1513-437: A separate compatible thallus being involved. These fungi are said to be homothallic, versus the normal heterothallic species with mating types. Others are secondarily homothallic, in that two compatible nuclei following meiosis migrate into each basidiospore, which is then dispersed as a pre-existing dikaryon. Often such species form only two spores per basidium, but that too varies. Following meiosis, mitotic divisions can occur in
1602-408: A substrate. These structures are called the sporodochium . This is a cushion of conidiophores created from a pseudoparenchymatous stroma in plant tissue. The pycnidium is a globose to flask-shaped parenchymatous structure, lined on its inner wall with conidiophores. The acervulus is a flat saucer shaped bed of conidiophores produced under a plant cuticle, which eventually erupt through
1691-557: A survival benefit for these fungi by promoting successful infection. A characteristic central feature of meiosis is recombination between homologous chromosomes. This process is associated with repair of DNA damage, particularly double-strand breaks . The ability of C. neoformans and M. maydis to undergo meiosis may contribute to their virulence by repairing the oxidative DNA damage caused by their host's release of reactive oxygen species . Many variations occur: some variations are self-compatible and spontaneously form dikaryons without
1780-551: A tetraploid nucleus which divided into four diploid nuclei by meiosis and then into eight haploid nuclei by a supposed process called brachymeiosis , but this hypothesis was disproven in the 1950s. From the fertilized ascogonium, dinucleate hyphae emerge in which each cell contains two nuclei. These hyphae are called ascogenous or fertile hyphae. They are supported by the vegetative mycelium containing uni– (or mono–) nucleate hyphae, which are sterile. The mycelium containing both sterile and fertile hyphae may grow into fruiting body,
1869-413: A typical Basidiomycota lifecycle the long lasting dikaryons periodically (seasonally or occasionally) produce basidia , the specialized usually club-shaped end cells, in which a pair of compatible nuclei fuse ( karyogamy ) to form a diploid cell. Meiosis follows shortly with the production of 4 haploid nuclei that migrate into 4 external, usually apical basidiospores. Variations occur, however. Typically
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#17327733168761958-423: A variety of stresses such as nutrient limitation. The sexual part of the life cycle commences when two hyphal structures mate . In the case of homothallic species, mating is enabled between hyphae of the same fungal clone , whereas in heterothallic species, the two hyphae must originate from fungal clones that differ genetically, i.e., those that are of a different mating type . Mating types are typical of
2047-417: Is a dictyospore . In staurospores ray-like arms radiate from a central body; in others ( helicospores ) the entire spore is wound up in a spiral like a spring. Very long worm-like spores with a length-to-diameter ratio of more than 15:1, are called scolecospores . Important characteristics of the anamorphs of the Ascomycota are conidiogenesis , which includes spore formation and dehiscence (separation from
2136-484: Is a tube-shaped vessel, a meiosporangium , which contains the sexual spores produced by meiosis and which are called ascospores . Apart from a few exceptions, such as Candida albicans , most ascomycetes are haploid , i.e., they contain one set of chromosomes per nucleus. During sexual reproduction there is a diploid phase, which commonly is very short, and meiosis restores the haploid state. The sexual cycle of one well-studied representative species of Ascomycota
2225-548: Is an example. By convention, the stages and spore states are numbered by Roman numerals . Typically, basidiospores infect host one, also known as the alternate or sexual host, and the mycelium forms pycnidia , which are miniature, flask-shaped, hollow, submicroscopic bodies embedded in the host tissue (such as a leaf). This stage, numbered "0", produces single-celled spores that ooze out in a sweet liquid and that act as nonmotile spermatia , and also protruding receptive hyphae . Insects and probably other vectors such as rain carry
2314-457: Is described in greater detail in Neurospora crassa . Also, the adaptive basis for the maintenance of sexual reproduction in the Ascomycota fungi was reviewed by Wallen and Perlin. They concluded that the most plausible reason for the maintenance of this capability is the benefit of repairing DNA damage by using recombination that occurs during meiosis . DNA damage can be caused by
2403-490: Is not known whether the fungus is parasitic , saprotrophic , or mutualistic . It is said to be edible. Ascomycetous Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota , forms the subkingdom Dikarya . Its members are commonly known as the sac fungi or ascomycetes . It is the largest phylum of Fungi, with over 64,000 species . The defining feature of this fungal group
2492-502: Is not yet placed in a subdivision, but recent genomic evidence suggests that it is a sister group of Agaricomycotina . The Pucciniomycotina include the rust fungi, the insect parasitic/symbiotic genus Septobasidium , a former group of smut fungi (in the Microbotryomycetes , which includes mirror yeasts), and a mixture of odd, infrequently seen, or seldom recognized fungi, often parasitic on plants. The eight classes in
2581-528: Is one of two large divisions that, together with the Ascomycota , constitute the subkingdom Dikarya (often referred to as the " higher fungi ") within the kingdom Fungi . Members are known as basidiomycetes . More specifically, Basidiomycota includes these groups: agarics , puffballs , stinkhorns , bracket fungi , other polypores , jelly fungi , boletes , chanterelles , earth stars , smuts , bunts , rusts , mirror yeasts , and Cryptococcus ,
2670-531: Is only found on Nothofagus (Southern Beech) in the Southern Hemisphere . Asexual reproduction is the dominant form of propagation in the Ascomycota, and is responsible for the rapid spread of these fungi into new areas. It occurs through vegetative reproductive spores, the conidia . The conidiospores commonly contain one nucleus and are products of mitotic cell divisions and thus are sometimes called mitospores, which are genetically identical to
2759-440: Is the " ascus " (from Ancient Greek ἀσκός ( askós ) 'sac, wineskin'), a microscopic sexual structure in which nonmotile spores , called ascospores , are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels , truffles , brewers' and bakers' yeast , dead man's fingers , and cup fungi . The fungal symbionts in
Neolecta - Misplaced Pages Continue
2848-423: Is the only order belonging to the class Neolectomycetes , which belongs to the subdivision Taphrinomycotina of the Ascomycota . Fruiting bodies take the shape of unbranched to lobed bright yellowish, orangish to pale yellow-green colored, club-shaped, smooth, fleshy columns up to about 7 centimetres (3 inches) tall. Neolecta fruitbodies consist of hyphae and a hymenium . The hymenium lacks paraphyses and
2937-463: Is usually inconspicuous because it is commonly embedded in the substrate, such as soil, or grows on or inside a living host, and only the ascoma may be seen when fruiting. Pigmentation , such as melanin in hyphal walls, along with prolific growth on surfaces can result in visible mold colonies; examples include Cladosporium species, which form black spots on bathroom caulking and other moist areas. Many ascomycetes cause food spoilage, and, therefore,
3026-416: The ascocarp , which may contain millions of fertile hyphae. An ascocarp is the fruiting body of the sexual phase in Ascomycota. There are five morphologically different types of ascocarp, namely: The sexual structures are formed in the fruiting layer of the ascocarp, the hymenium . At one end of ascogenous hyphae, characteristic U-shaped hooks develop, which curve back opposite to the growth direction of
3115-402: The ascogonium , and merges with a gametangium (the antheridium ) of the other fungal isolate. The nuclei in the antheridium then migrate into the ascogonium, and plasmogamy —the mixing of the cytoplasm —occurs. Unlike in animals and plants, plasmogamy is not immediately followed by the merging of the nuclei (called karyogamy ). Instead, the nuclei from the two hyphae form pairs, initiating
3204-473: The Ascomycota and has been called a living fossil . The genome of N. irregularis has been sequenced. Sequence analysis has revealed that rudimentary multicellularity is deeply rooted in the Ascomycota. Neolecta is found in Asia, Northern Europe, North America, and southern Brazil. The species all live in association with trees, and at least one, N. vitellina , grows from rootlets of its host, but it
3293-762: The Gasteromycetes (another obsolete class that included species mostly lacking hymenia and mostly forming spores in enclosed fruitbodies ), as well as most of the jelly fungi . This sub-phyla also includes the "classic" mushrooms, polypores, corals, chanterelles, crusts, puffballs and stinkhorns. The three classes in the Agaricomycotina are the Agaricomycetes , the Dacrymycetes , and the Tremellomycetes . The class Wallemiomycetes
3382-451: The Rust ( Pucciniales )) tend to have mutually indistinguishable, compatible haploids which are usually mycelia being composed of filamentous hyphae . Typically haploid Basidiomycota mycelia fuse via plasmogamy and then the compatible nuclei migrate into each other's mycelia and pair up with the resident nuclei. Karyogamy is delayed, so that the compatible nuclei remain in pairs, called
3471-450: The asci lack croziers , which makes the genus distinctive among other earth-tongues. Neolecta vitellina forms masses of conidia by budding, hinting at the possibility that it also produces a yeast state. Species of the genus may resemble those of Clavulinopsis and Spathularia . Neolecta does not have any close relatives. Phylogenetically, it clusters weakly with a bizarre group of basal Ascomycota including: Taphrina ,
3560-456: The conidia . The asexual, non-motile haploid spores of a fungus, which are named after the Greek word for dust (conia), are hence also known as conidiospores . The conidiospores commonly contain one nucleus and are products of mitotic cell divisions and thus are sometimes called mitospores , which are genetically identical to the mycelium from which they originate. They are typically formed at
3649-627: The detritivores (animals that feed on decomposing material) to obtain their nutrients. Ascomycetes, along with other fungi, can break down large molecules such as cellulose or lignin , and thus have important roles in nutrient cycling such as the carbon cycle . The fruiting bodies of the Ascomycota provide food for many animals ranging from insects and slugs and snails ( Gastropoda ) to rodents and larger mammals such as deer and wild boars . Many ascomycetes also form symbiotic relationships with other organisms, including plants and animals. Probably since early in their evolutionary history,
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3738-401: The dikaryophase of the sexual cycle, during which time the pairs of nuclei synchronously divide. Fusion of the paired nuclei leads to mixing of the genetic material and recombination and is followed by meiosis . A similar sexual cycle is present in the red algae (Rhodophyta). A discarded hypothesis held that a second karyogamy event occurred in the ascogonium prior to ascogeny, resulting in
3827-685: The ergot fungi, black knot , and the powdery mildews . The members of the genus Cordyceps are entomopathogenic fungi , meaning that they parasitise and kill insects. Other entomopathogenic ascomycetes have been used successfully in biological pest control , such as Beauveria . Several species of ascomycetes are biological model organisms in laboratory research. Most famously, Neurospora crassa , several species of yeasts , and Aspergillus species are used in many genetics and cell biology studies. Ascomycetes are 'spore shooters'. They are fungi which produce microscopic spores inside special, elongated cells or sacs, known as 'asci', which give
3916-425: The photoautotrophic algal partner generates metabolic energy through photosynthesis, the fungus offers a stable, supportive matrix and protects cells from radiation and dehydration. Around 42% of the Ascomycota (about 18,000 species) form lichens, and almost all the fungal partners of lichens belong to the Ascomycota. Basidiomycota Basidiomycota ( / b ə ˌ s ɪ d i . oʊ m aɪ ˈ k oʊ t ə / )
4005-542: The Ascomycota have formed symbiotic associations with green algae ( Chlorophyta ), and other types of algae and cyanobacteria . These mutualistic associations are commonly known as lichens , and can grow and persist in terrestrial regions of the earth that are inhospitable to other organisms and characterized by extremes in temperature and humidity, including the Arctic , the Antarctic , deserts , and mountaintops. While
4094-410: The Ascomycota. The most frequent types are the single-celled spores, which are designated amerospores . If the spore is divided into two by a cross-wall ( septum ), it is called a didymospore . When there are two or more cross-walls, the classification depends on spore shape. If the septae are transversal , like the rungs of a ladder, it is a phragmospore , and if they possess a net-like structure it
4183-649: The Ascomycota. These include the following sexual ( teleomorphic ) groups, defined by the structures of their sexual fruiting bodies : the Discomycetes , which included all species forming apothecia ; the Pyrenomycetes , which included all sac fungi that formed perithecia or pseudothecia , or any structure resembling these morphological structures; and the Plectomycetes, which included those species that form cleistothecia . Hemiascomycetes included
4272-491: The Basidiomycota that are 1) poorly known, 2) have not been subjected to DNA analysis, or 3) if analysed phylogenetically do not group with as yet named or identified families, and have not been assigned to a specific family (i.e., they are incertae sedis with respect to familial placement). These include: Unlike animals and plants which have readily recognizable male and female counterparts, Basidiomycota (except for
4361-520: The Basidiomycota were divided into two classes, now obsolete: Nonetheless these former concepts continue to be used as two types of growth habit groupings, the "mushrooms" (e.g. Schizophyllum commune ) and the non-mushrooms (e.g. Mycosarcoma maydis ). The Agaricomycotina include what had previously been called the Hymenomycetes (an obsolete morphological based class of Basidiomycota that formed hymenial layers on their fruitbodies ),
4450-874: The Basidiomycota. As now classified, the subphyla join and also cut across various obsolete taxonomic groups (see below) previously commonly used to describe Basidiomycota. According to a 2008 estimate, Basidiomycota comprise three subphyla (including six unassigned classes) 16 classes, 52 orders, 177 families, 1,589 genera, and 31,515 species. Wijayawardene et al. 2020 produced an update that recognized 19 classes ( Agaricomycetes , Agaricostilbomycetes , Atractiellomycetes , Bartheletiomycetes , Classiculomycetes , Cryptomycocolacomycetes , Cystobasidiomycetes , Dacrymycetes , Exobasidiomycetes , Malasseziomycetes , Microbotryomycetes , Mixiomycetes , Monilielliomycetes , Pucciniomycetes , Spiculogloeomycetes , Tremellomycetes , Tritirachiomycetes , Ustilaginomycetes and Wallemiomycetes ) with multiple orders and genera. Traditionally,
4539-864: The Pucciniomycotina are Agaricostilbomycetes , Atractiellomycetes , Classiculomycetes , Cryptomycocolacomycetes , Cystobasidiomycetes , Microbotryomycetes , Mixiomycetes , and Pucciniomycetes . The Ustilaginomycotina are most (but not all) of the former smut fungi and the Exobasidiales . The classes of the Ustilaginomycotina are the Exobasidiomycetes , the Entorrhizomycetes, and the Ustilaginomycetes . There are several genera classified in
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#17327733168764628-482: The absence of dikaryon formation, diploid nuclei, and meiosis. A rare few number of taxa have extended diploid lifecycles, but can be common species. Examples exist in the mushroom genera Armillaria and Xerula , both in the Physalacriaceae . Occasionally, basidiospores are not formed and parts of the "basidia" act as the dispersal agents, e.g. the peculiar mycoparasitic jelly fungus, Tetragoniomyces or
4717-453: The atmosphere and freshwater environments, as well as ocean beaches and tidal zones. The distribution of species is variable; while some are found on all continents, others, as for example the white truffle Tuber magnatum , only occur in isolated locations in Italy and Eastern Europe. The distribution of plant-parasitic species is often restricted by host distributions; for example, Cyttaria
4806-533: The basidiospores are ballistic , hence they are sometimes also called ballistospores . In most species, the basidiospores disperse and each can start a new haploid mycelium, continuing the lifecycle. Basidia are microscopic but they are often produced on or in multicelled large fructifications called basidiocarps or basidiomes, or fruitbodies , variously called mushrooms, puffballs , etc. Ballistic basidiospores are formed on sterigmata which are tapered spine-like projections on basidia, and are typically curved, like
4895-633: The basidium. Multiple numbers of basidiospores can result, including odd numbers via degeneration of nuclei, or pairing up of nuclei, or lack of migration of nuclei. For example, the chanterelle genus Craterellus often has six-spored basidia, while some corticioid Sistotrema species can have two-, four-, six-, or eight-spored basidia, and the cultivated button mushroom, Agaricus bisporus . can have one-, two-, three- or four-spored basidia under some circumstances. Occasionally, monokaryons of some taxa can form morphologically fully formed basidiomes and anatomically correct basidia and ballistic basidiospores in
4984-755: The cell. Many species live on dead plant material such as leaves, twigs, or logs. Several species colonize plants, animals, or other fungi as parasites or mutualistic symbionts and derive all their metabolic energy in form of nutrients from the tissues of their hosts. Owing to their long evolutionary history, the Ascomycota have evolved the capacity to break down almost every organic substance. Unlike most organisms, they are able to use their own enzymes to digest plant biopolymers such as cellulose or lignin . Collagen , an abundant structural protein in animals, and keratin —a protein that forms hair and nails—, can also serve as food sources. Unusual examples include Aureobasidium pullulans , which feeds on wall paint, and
5073-421: The core expression program of meiosis has been conserved in these fungi for over half a billion years of evolution since these species diverged. Cryptococcus neoformans and Mycosarcoma maydis are examples of pathogenic basidiomycota. Such pathogens must be able to overcome the oxidative defenses of their respective hosts in order to produce a successful infection. The ability to undergo meiosis may provide
5162-417: The cuticle for dispersal. Asexual reproduction process in ascomycetes also involves the budding which we clearly observe in yeast. This is termed a "blastic process". It involves the blowing out or blebbing of the hyphal tip wall. The blastic process can involve all wall layers, or there can be a new cell wall synthesized which is extruded from within the old wall. The initial events of budding can be seen as
5251-605: The development of a ring of chitin around the point where the bud is about to appear. This reinforces and stabilizes the cell wall. Enzymatic activity and turgor pressure act to weaken and extrude the cell wall. New cell wall material is incorporated during this phase. Cell contents are forced into the progeny cell, and as the final phase of mitosis ends a cell plate, the point at which a new cell wall will grow inwards from, forms. There are three subphyla that are described and accepted: Several outdated taxon names—based on morphological features—are still occasionally used for species of
5340-517: The elongated haploid basidiospores form apically, often in compatible pairs that fuse centrally resulting in H-shaped diaspores which are by then dikaryotic. Dikaryotic conidia may then form. Eventually the host is infected by infectious hyphae. Teliospores form in host tissue. Many variations on these general themes occur. Smuts with both a yeast phase and an infectious hyphal state are examples of dimorphic Basidiomycota. In plant parasitic taxa,
5429-407: The ends of specialized hyphae, the conidiophores. Depending on the species they may be dispersed by wind or water, or by animals. Conidiophores may simply branch off from the mycelia or they may be formed in fruiting bodies. The hypha that creates the sporing (conidiating) tip can be very similar to the normal hyphal tip, or it can be differentiated. The most common differentiation is the formation of
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#17327733168765518-865: The entire "basidium" acts as a "spore", e.g. in some false puffballs ( Scleroderma ). In the human pathogenic genus Cryptococcus , four nuclei following meiosis remain in the basidium, but continually divide mitotically, each nucleus migrating into synchronously forming nonballistic basidiospores that are then pushed upwards by another set forming below them, resulting in four parallel chains of dry "basidiospores". Other variations occur: some as standard lifecycles (that themselves have variations within variations) within specific orders. Rusts ( Pucciniales , previously known as Uredinales ) at their greatest complexity, produce five different types of spores on two different host plants in two unrelated host families. Such rusts are heteroecious (requiring two hosts) and macrocyclic (producing all five spores types). Wheat stem rust
5607-423: The formation of a distinctive anatomical feature (the clamp connection ), cell wall components, and definitively by phylogenetic molecular analysis of DNA sequence data. A 2007 classification, adopted by a coalition of 67 mycologists recognized three subphyla (Pucciniomycotina, Ustilaginomycotina, Agaricomycotina) and two other class level taxa ( Wallemiomycetes , Entorrhizomycetes ) outside of these, among
5696-663: The fungal symbiont directly obtains products of photosynthesis . In common with many basidiomycetes and Glomeromycota , some ascomycetes form symbioses with plants by colonizing the roots to form mycorrhizal associations. The Ascomycota also represents several carnivorous fungi , which have developed hyphal traps to capture small protists such as amoebae , as well as roundworms ( Nematoda ), rotifers , tardigrades , and small arthropods such as springtails ( Collembola ). The Ascomycota are represented in all land ecosystems worldwide, occurring on all continents including Antarctica . Spores and hyphal fragments are dispersed through
5785-428: The fungi and correspond roughly to the sexes in plants and animals; however one species may have more than two mating types, resulting in sometimes complex vegetative incompatibility systems. The adaptive function of mating type is discussed in Neurospora crassa . Gametangia are sexual structures formed from hyphae, and are the generative cells. A very fine hypha, called trichogyne emerges from one gametangium,
5874-517: The group its name. Asexual reproduction is the dominant form of propagation in the Ascomycota, and is responsible for the rapid spread of these fungi into new areas. Asexual reproduction of ascomycetes is very diverse from both structural and functional points of view. The most important and general is production of conidia, but chlamydospores are also frequently produced. Furthermore, Ascomycota also reproduce asexually through budding. Asexual reproduction may occur through vegetative reproductive spores,
5963-523: The hook with one nucleus, one at the basal of the original hypha that contains one nucleus, and one that separates the U-shaped part, which contains the other two nuclei. Fusion of the nuclei (karyogamy) takes place in the U-shaped cells in the hymenium, and results in the formation of a diploid zygote . The zygote grows into the ascus , an elongated tube-shaped or cylinder-shaped capsule. Meiosis then gives rise to four haploid nuclei, usually followed by
6052-702: The horns of a bull. In some Basidiomycota the spores are not ballistic, and the sterigmata may be straight, reduced to stubs, or absent. The basidiospores of these non-ballistosporic basidia may either bud off, or be released via dissolution or disintegration of the basidia. In summary, meiosis takes place in a diploid basidium. Each one of the four haploid nuclei migrates into its own basidiospore. The basidiospores are ballistically discharged and start new haploid mycelia called monokaryons. There are no males or females, rather there are compatible thalli with multiple compatibility factors. Plasmogamy between compatible individuals leads to delayed karyogamy leading to establishment of
6141-542: The human pathogenic yeast. Basidiomycota are filamentous fungi composed of hyphae (except for basidiomycota-yeast ) and reproduce sexually via the formation of specialized club-shaped end cells called basidia that normally bear external meiospores (usually four). These specialized spores are called basidiospores . However, some Basidiomycota are obligate asexual reproducers. Basidiomycota that reproduce asexually (discussed below) can typically be recognized as members of this division by gross similarity to others, by
6230-446: The hyphae, called " septa ", are the internal boundaries of individual cells (or compartments). The cell wall and septa give stability and rigidity to the hyphae and may prevent loss of cytoplasm in case of local damage to cell wall and cell membrane . The septa commonly have a small opening in the center, which functions as a cytoplasmic connection between adjacent cells, also sometimes allowing cell-to-cell movement of nuclei within
6319-416: The hyphae. The two nuclei contained in the apical part of each hypha divide in such a way that the threads of their mitotic spindles run parallel, creating two pairs of genetically different nuclei. One daughter nucleus migrates close to the hook, while the other daughter nucleus locates to the basal part of the hypha. The formation of two parallel cross-walls then divides the hypha into three sections: one at
6408-577: The infection process, rather it remains dormant for a period and then germinates to form basidia (stage "IV"), sometimes called a promycelium . In the Pucciniales, the basidia are cylindrical and become 3- septate after meiosis, with each of the 4 cells bearing one basidiospore each. The basidiospores disperse and start the infection process on host 1 again. Autoecious rusts complete their life-cycles on one host instead of two, and microcyclic rusts cut out one or more stages. The characteristic part of
6497-612: The kerosene fungus Amorphotheca resinae , which feeds on aircraft fuel (causing occasional problems for the airline industry), and may sometimes block fuel pipes. Other species can resist high osmotic stress and grow, for example, on salted fish, and a few ascomycetes are aquatic. The Ascomycota is characterized by a high degree of specialization; for instance, certain species of Laboulbeniales attack only one particular leg of one particular insect species. Many Ascomycota engage in symbiotic relationships such as in lichens—symbiotic associations with green algae or cyanobacteria —in which
6586-405: The life-cycle of smuts is the thick-walled, often darkly pigmented, ornate, teliospore that serves to survive harsh conditions such as overwintering and also serves to help disperse the fungus as dry diaspores . The teliospores are initially dikaryotic but become diploid via karyogamy. Meiosis takes place at the time of germination. A promycelium is formed that consists of a short hypha (equated to
6675-1159: The majority of lichens (loosely termed "ascolichens") such as Cladonia belong to the Ascomycota. Ascomycota is a monophyletic group (containing all of the descendants of a common ancestor). Previously placed in the Basidiomycota along with asexual species from other fungal taxa, asexual (or anamorphic ) ascomycetes are now identified and classified based on morphological or physiological similarities to ascus-bearing taxa , and by phylogenetic analyses of DNA sequences. Ascomycetes are of particular use to humans as sources of medicinally important compounds such as antibiotics , as well as for fermenting bread, alcoholic beverages, and cheese. Examples of ascomycetes include Penicillium species on cheeses and those producing antibiotics for treating bacterial infectious diseases . Many ascomycetes are pathogens , both of animals, including humans, and of plants. Examples of ascomycetes that can cause infections in humans include Candida albicans , Aspergillus niger and several tens of species that cause skin infections . The many plant-pathogenic ascomycetes include apple scab , rice blast ,
6764-403: The mushroom cap, and the meiotic prophase stage is prolonged. Burns et al. studied the expression of genes involved in the 15-hour meiotic process, and found that the pattern of gene expression of C. cinerea was similar to two other fungal species, the yeasts Saccharomyces cerevisiae and Schizosaccharomyces pombe . These similarities in the patterns of expression led to the conclusion that
6853-402: The mycelium from which they originate. They are typically formed at the ends of specialized hyphae , the conidiophores . Depending on the species they may be dispersed by wind or water, or by animals. Different types of asexual spores can be identified by colour, shape, and how they are released as individual spores. Spore types can be used as taxonomic characters in the classification within
6942-436: The original parent nuclei. Alternatively, nuclei may lose some chromosomes, resulting in aneuploid cells. Candida albicans (class Saccharomycetes) is an example of a fungus that has a parasexual cycle (see Candida albicans and Parasexual cycle ). Sexual reproduction in the Ascomycota leads to the formation of the ascus , the structure that defines this fungal group and distinguishes it from other fungal phyla. The ascus
7031-413: The parent structure). Conidiogenesis corresponds to Embryology in animals and plants and can be divided into two fundamental forms of development: blastic conidiogenesis, where the spore is already evident before it separates from the conidiogenic hypha, and thallic conidiogenesis, during which a cross-wall forms and the newly created cell develops into a spore. The spores may or may not be generated in
7120-472: The pellicles or moldy layers that develop on jams, juices, and other foods are the mycelia of these species or occasionally Mucoromycotina and almost never Basidiomycota . Sooty molds that develop on plants, especially in the tropics are the thalli of many species. Large masses of yeast cells, asci or ascus-like cells, or conidia can also form macroscopic structures. For example. Pneumocystis species can colonize lung cavities (visible in x-rays), causing
7209-705: The pleiomorphic rusts are examples of fungi with anamorphs , which are the asexual stages. Some Basidiomycota are only known as anamorphs. Many are called basidiomycetous yeasts, which differentiates them from ascomycetous yeasts in the Ascomycota . Aside from yeast anamorphs and uredinia, aecia, and pycnidia, some Basidiomycota form other distinctive anamorphs as parts of their life cycles. Examples are Collybia tuberosa with its apple-seed-shaped and coloured sclerotium , Dendrocollybia racemosa with its sclerotium and its Tilachlidiopsis racemosa conidia, Armillaria with their rhizomorphs , Hohenbuehelia with their Nematoctonus nematode infectious, state and
7298-436: The primary host a repeating spore stage is formed, numbered "II", the urediospores in dry pustules called uredinia . Urediospores are dikaryotic and can infect the same host that produced them. They repeatedly infect this host over the growing season. At the end of the season, a fourth spore type, the teliospore , is formed. It is thicker-walled and serves to overwinter or to survive other harsh conditions. It does not continue
7387-656: The process of heterokaryosis, caused by merging of two hyphae belonging to different individuals, by a process called anastomosis , followed by a series of events resulting in genetically different cell nuclei in the mycelium . The merging of nuclei is not followed by meiotic events , such as gamete formation and results in an increased number of chromosomes per nuclei. Mitotic crossover may enable recombination , i.e., an exchange of genetic material between homologous chromosomes . The chromosome number may then be restored to its haploid state by nuclear division , with each daughter nuclei being genetically different from
7476-766: The same species. Species of the Deuteromycota were classified as Coelomycetes if they produced their conidia in minute flask- or saucer-shaped conidiomata, known technically as pycnidia and acervuli . The Hyphomycetes were those species where the conidiophores ( i.e. , the hyphal structures that carry conidia-forming cells at the end) are free or loosely organized. They are mostly isolated but sometimes also appear as bundles of cells aligned in parallel (described as synnematal ) or as cushion-shaped masses (described as sporodochial ). Most species grow as filamentous, microscopic structures called hyphae or as budding single cells (yeasts). Many interconnected hyphae form
7565-495: The saprotrophic phase is normally the yeast while the infectious stage is hyphal. However, there are examples of animal and human parasites where the species are dimorphic but it is the yeast-like state that is infectious. The genus Filobasidiella forms basidia on hyphae but the main infectious stage is more commonly known by the anamorphic yeast name Cryptococcus , e.g. Cryptococcus neoformans and Cryptococcus gattii . The dimorphic Basidiomycota with yeast stages and
7654-448: The spermatia from spermagonium to spermagonium, cross inoculating the mating types. Neither thallus is male or female. Once crossed, the dikaryons are established and a second spore stage is formed, numbered "I" and called aecia , which form dikaryotic aeciospores in dry chains in inverted cup-shaped bodies embedded in host tissue. These aeciospores then infect the second host, known as the primary or asexual host (in macrocyclic rusts). On
7743-514: The spores reach a suitable substrate, they germinate, form new hyphae, which restarts the fungal life cycle. The form of the ascus is important for classification and is divided into four basic types: unitunicate-operculate, unitunicate-inoperculate, bitunicate, or prototunicate. See the article on asci for further details. The Ascomycota fulfil a central role in most land-based ecosystems . They are important decomposers , breaking down organic materials, such as dead leaves and animals, and helping
7832-409: The thallus of the fungus defines the shape of the symbiotic colony. Some dimorphic species, such as Candida albicans , can switch between growth as single cells and as filamentous, multicellular hyphae. Other species are pleomorphic , exhibiting asexual (anamorphic) as well as a sexual (teleomorphic) growth forms. Except for lichens, the non-reproductive (vegetative) mycelium of most ascomycetes
7921-857: The yeasts and yeast-like fungi that have now been placed into the Saccharomycotina or Taphrinomycotina , while the Euascomycetes included the remaining species of the Ascomycota, which are now in the Pezizomycotina , and the Neolecta , which are in the Taphrinomycotina. Some ascomycetes do not reproduce sexually or are not known to produce asci and are therefore anamorphic species. Those anamorphs that produce conidia (mitospores) were previously described as mitosporic Ascomycota . Some taxonomists placed this group into
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