np.

真菌、菌類、真菌類

関: filamentous fungi、fungal、fungus、microbial、mold、myco

WordNet

fit tightly, follow the contours of; "The dress molds her beautiful figure"
become moldy; spoil due to humidity; "The furniture molded in the old house" (同)mildew
a fungus that produces a superficial growth on various kinds of damp or decaying organic matter (同)mould
loose soil rich in organic matter (同)mould
a distinctive nature, character, or type; "a leader in the mold of her predecessors" (同)mould
container into which liquid is poured to create a given shape when it hardens (同)mould, cast
sculpture produced by molding (同)mould, molding, moulding, modeling, clay sculpture
a dish or dessert that is formed in or on a mold; "a lobster mold"; "a gelatin dessert made in a mold" (同)mould
an organism of the kingdom Fungi lacking chlorophyll and feeding on organic matter; ranging from unicellular or multicellular organisms to spore-bearing syncytia
of or involving or caused by or being microbes; "microbial warfare" (同)microbic
of or relating to fungi (同)fungous
activities that are enjoyable or amusing; "I do it for the fun of it"; "he is fun to have around" (同)merriment, playfulness
violent and excited activity; "she asked for money and then the fun began"; "they began to fight like fun"
verbal wit or mockery (often at anothers expense but not to be taken seriously); "he became a figure of fun"; "he said it in sport" (同)play, sport
the taxonomic kingdom including yeast, molds, smuts, mushrooms, and toadstools; distinct from the green plants (同)kingdom_Fungi, fungus kingdom

PrepTutorEJDIC

菌(葉緑素が無く,他の生物の有機物に依存しているキノコ,カビなど)
『楽しみ』,慰み / 戯れ,ふざけ / 愉快な気持ちにさせるもの(人)

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2013/11/26 17:14:23」(JST)

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A fungus (/ˈfʌŋɡəs/; plural: fungi^[3] or funguses^[4]) is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds (British English: moulds), as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, protists and bacteria. One major difference is that fungal cells have cell walls that contain chitin, unlike the cell walls of plants and some protists, which contain cellulose, and unlike the cell walls of bacteria. These and other differences show that the fungi form a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (is a monophyletic group). This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης, mukēs, meaning "fungus"). Mycology has often been regarded as a branch of botany, even though it is a separate kingdom in biological taxonomy. Genetic studies have shown that fungi are more closely related to animals than to plants.

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil, on dead matter, and as symbionts of plants, animals, or other fungi. They may become noticeable when fruiting, either as mushrooms or molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange. They have long been used as a direct source of food, such as mushrooms and truffles, as a leavening agent for bread, and in fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g. rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from single-celled aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of Kingdom Fungi, which has been estimated at 1.5 million to 5 million species, with about 5% of these having been formally classified. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christian Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the last decade have helped reshape the classification of Kingdom Fungi, which is divided into one subkingdom, seven phyla, and ten subphyla.

A group of all the fungi present in a particular area or geographic region is known as mycobiota (plural noun, no singular), e.g. "the mycobiota of Ireland".^[5]

1 Etymology
2 Characteristics
3 Diversity
4 Morphology
- 4.1 Microscopic structures
- 4.2 Macroscopic structures
5 Growth and physiology
6 Reproduction
- 6.1 Asexual reproduction
- 6.2 Sexual reproduction
- 6.3 Spore dispersal
- 6.4 Other sexual processes
7 Evolution
8 Taxonomy
- 8.1 Taxonomic groups
- 8.2 Fungus-like organisms
9 Ecology
- 9.1 Symbiosis
  - 9.1.1 With plants
  - 9.1.2 With algae and cyanobacteria
  - 9.1.3 With insects
  - 9.1.4 As pathogens and parasites
10 Pathogenic mechanisms
11 Human use
- 11.1 Drugs
- 11.2 Cultured foods
- 11.3 Medicinal use
- 11.4 Edible and poisonous species
- 11.5 Pest control
- 11.6 Bioremediation
- 11.7 Model organisms
- 11.8 Others
12 Mycotoxins
13 Mycology
- 13.1 History
14 See also
15 References
- 15.1 Cited literature
16 External links

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny.^[6] This in turn is derived from the Greek word sphongos/σφογγος ("sponge"), which refers to the macroscopic structures and morphology of mushrooms and molds;^[7] the root is also used in other languages, such as the German Schwamm ("sponge") and Schimmel ("mold").^[8] The use of the word mycology, which is derived from the Greek mykes/μύκης (mushroom) and logos/λόγος (discourse),^[9] to denote the scientific study of fungi is thought to have originated in 1836 with English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5.^[7]

Characteristics

Fungal Hyphae Cells

1- Hyphal wall 2- Septum 3- Mitochondrion 4- Vacuole 5- Ergosterol crystal 6- Ribosome 7- Nucleus 8- Endoplasmic reticulum 9- Lipid body 10- Plasma membrane 11- Spitzenkörper 12- Golgi apparatus

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the Plant Kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Like plants, fungi often grow in soil, and in the case of mushrooms form conspicuous fruiting bodies, which sometimes bear resemblance to plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago.^[10]^[11] Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

Shared features:

With other eukaryotes: As other eukaryotes, fungal cells contain membrane-bound nuclei with chromosomes that contain DNA with noncoding regions called introns and coding regions called exons. In addition, fungi possess membrane-bound cytoplasmic organelles such as mitochondria, sterol-containing membranes, and ribosomes of the 80S type.^[12] They have a characteristic range of soluble carbohydrates and storage compounds, including sugar alcohols (e.g., mannitol), disaccharides, (e.g., trehalose), and polysaccharides (e.g., glycogen, which is also found in animals^[13]).
With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.^[14]
With plants: Fungi possess a cell wall^[15] and vacuoles.^[16] They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.^[17]
With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.^[18]^[19]
The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures and are called hyphae, which may contain multiple nuclei and extend at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called Spitzenkörper.^[20] Both fungi and oomycetes grow as filamentous hyphal cells.^[21] In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells.^[13]
In common with some plant and animal species, more than 60 fungal species display the phenomenon of bioluminescence.^[22]

Unique features:

Some species grow as single-celled yeasts that reproduce by budding or binary fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.^[23]
The fungal cell wall is composed of glucans and chitin; while the former compounds are also found in plants and the latter in the exoskeleton of arthropods,^[24]^[25] fungi are the only organisms that combine these two structural molecules in their cell wall. In contrast to plants and the oomycetes, fungal cell walls do not contain cellulose.^[26]

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome these limitations, some fungi, such as Armillaria, form rhizomorphs,^[27] that resemble and perform functions similar to the roots of plants. Another characteristic shared with plants includes a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks.^[28] However, plants have an additional terpene pathway in their chloroplasts, a structure fungi do not possess.^[29] Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants.^[28] Many of the plant and fungal enzymes that make these compounds differ from each other in sequence and other characteristics, which indicates separate origins and evolution of these enzymes in the fungi and plants.^[28]^[30]

Diversity

Bracket fungi on dead log

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations^[31] or ionizing radiation,^[32] as well as in deep sea sediments.^[33] Some can survive the intense UV and cosmic radiation encountered during space travel.^[34] Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungus Batrachochytrium dendrobatidis, a parasite that has been responsible for a worldwide decline in amphibian populations. This organism spends part of its life cycle as a motile zoospore, enabling it to propel itself through water and enter its amphibian host.^[35] Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.^[36]

Around 100,000 species of fungi have been formally described by taxonomists,^[37] but the global biodiversity of the fungus kingdom is not fully understood.^[38] On the basis of observations of the ratio of the number of fungal species to the number of plant species in selected environments, the fungal kingdom has been estimated to contain about 1.5 million species;^[39] a recent (2011) estimate suggests there may be over 5 million species.^[40] In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy.^[41] Species may also be distinguished by their biochemical and physiological characteristics, such as their ability to metabolize certain biochemicals, or their reaction to chemical tests. The biological species concept discriminates species based on their ability to mating in fungi. The application of molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.^[42]

Morphology

Microscopic structures

An environmental isolate of Penicillium

1. hypha 2. conidiophore 3. phialide 4. conidia 5. septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips bifurcate (fork) giving rise to two parallel-growing hyphae.^[43] The combination of apical growth and branching/forking leads to the development of a mycelium, an interconnected network of hyphae.^[23] Hyphae can be either septate or coenocytic: septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized.^[44] Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in the fungi of the phylum Basidiomycota.^[45] Coenocytic hyphae are essentially multinucleate supercells.^[46]

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla, and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.^[47]

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella.^[48] Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.^[49]

Macroscopic structures

Armillaria solidipes

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and on spoiled food, where they are commonly called molds. Mycelia grown on solid agar media in laboratory petri dishes are usually referred to as colonies. These colonies can exhibit growth shapes and colors (due to spores or pigmentation) that can be used as diagnostic features in the identification of species or groups.^[50] Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.^[51]

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruiting body that holds the hymenium, a layer of tissue containing the spore-bearing cells.^[52] The fruiting bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

Growth and physiology

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios.^[53] Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues.^[54] They can exert large penetrative mechanical forces; for example, the plant pathogen Magnaporthe grisea forms a structure called an appressorium that evolved to puncture plant tissues.^[55] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).^[55] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.^[56]

The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing osmolytes such as glycerol.^[57] Morphological adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic molecules—such as polysaccharides, proteins, lipids, and other organic substrates—into smaller molecules that may then be absorbed as nutrients.^[58]^[59]^[60] The vast majority of filamentous fungi grow in a polar fashion—i.e., by extension into one direction—by elongation at the tip (apex) of the hypha.^[61] Alternative forms of fungal growth include intercalary extension (i.e., by longitudinal expansion of hyphal compartments that are below the apex) as in the case of some endophytic fungi,^[62] or growth by volume expansion during the development of mushroom stipes and other large organs.^[63] Growth of fungi as multicellular structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants^[64]—has several functions, including the development of fruiting bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication.^[65]

The fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol.^[66]^[67] For some species it has been shown that the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation; however, this form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not known.^[32] The authors speculate that this process might bear similarity to CO₂ fixation via visible light, but instead utilizing ionizing radiation as a source of energy.^[68]

Reproduction

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms.^[69] It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph and the anamorph.^[70] Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

Asexual reproduction

Asexual reproduction via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegatative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction.^[71] The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species which lack an observable sexual cycle.^[72]

Sexual reproduction

Spore dispersal

Both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

The bird's nest fungus Cyathus stercoreus

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection.^[82] For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air.^[83] The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g;^[84] the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below.^[85] Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies.^[86] Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.^[87]

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells.^[88] The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization^[89] and is likely required for hybridization between species, which has been associated with major events in fungal evolution.^[90]

Evolution

Main article: Evolution of fungi

In contrast to plants and animals, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal fruiting bodies, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi.^[91] Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy.^[92] Compression fossils are studied by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.^[93]

The earliest fossils possessing features typical of fungi date to the Proterozoic eon, some 1,430 million years ago (Ma); these multicellular benthic organisms had filamentous structures with septa, and were capable of anastomosis.^[94] More recent studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups.^[95] For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores.^[96] The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization.^[97] Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.^[98]

The fungi probably colonized the land during the Cambrian (542–488.3 Ma), long before land plants.^[99] Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants.^[100] Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they are abundant in the Rhynie chert, mostly as Zygomycota and Chytridiomycota.^[99]^[101]^[102] At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged,^[103] and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).^[104]

Lichen-like fossils have been found in the Doushantuo Formation in southern China dating back to 635–551 Ma.^[105] Lichens were a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 400 Ma;^[106] this date corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert.^[107] The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian.^[108] Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the mid-Cretaceous, 90 Ma.^[109]^[110]

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period.^[111] However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess,^[112] the spike did not appear worldwide,^[113]^[114] and in many places it did not fall on the Permian–Triassic boundary.^[115]

Taxonomy

Unikonta

Amoebozoa

Opisthokonta

	Animalia

	Choanozoa

Nucleariids

Rozellida

Fungi^[42]

Microsporidia

	Chytridiomycota

	Neocallimastigomycota

Blastocladiomycota

Zoopagomycotina

Kickxellomycotina

Entomophthoromycotina

Mucoromycotina

Glomeromycota

Dikarya

	Ascomycota

	Basidiomycota

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts.^[116] Analyses using molecular phylogenetics support a monophyletic origin of the Fungi.^[42] The taxonomy of the Fungi is in a state of constant flux, especially due to recent research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.^[117]

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature.^[42]^[118] Fungal species can also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and ITIS list current names of fungal species (with cross-references to older synonyms).

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy.^[42] It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya. The below cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms. The lengths of the branches in this tree are not proportional to evolutionary distances.

Taxonomic groups

Fungus-like organisms

Because of similarities in morphology and lifestyle, the slime molds (myxomycetes) and water molds (oomycetes) were formerly classified in the kingdom Fungi. Unlike true fungi the cell walls of these organisms contain cellulose and lack chitin. Myxomycetes are unikonts like fungi, but are grouped in the Amoebozoa. Oomycetes are diploid bikonts, grouped in the Chromalveolate kingdom. Neither water molds nor slime molds are closely related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology textbooks and primary research literature.^[130]

The Rozellida clade, including the "chytrid" Rozella, is a genetically disparate group known mostly from environmental DNA sequences that is a sister group to fungi. Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi.

The nucleariids, currently grouped in the Choanozoa, may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom.^[131]

Ecology

A pin mould in the process of decomposing a peach

Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Along with bacteria, fungi are the major decomposers in most terrestrial (and some aquatic) ecosystems, and therefore play a critical role in biogeochemical cycles^[132] and in many food webs. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms.^[133]^[134]

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all Kingdoms.^[135]^[136]^[137] These interactions can be mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.^[138]^[139]^[140]

With plants

Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant–fungus associations and is of significant importance for plant growth and persistence in many ecosystems; over 90% of all plant species engage in mycorrhizal relationships with fungi and are dependent upon this relationship for survival.^[141]

The dark filaments are hyphae of the endophytic fungus Neotyphodium coenophialum in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating to at least 400 million years ago.^[122] It often increases the plant's uptake of inorganic compounds, such as nitrate and phosphate from soils having low concentrations of these key plant nutrients.^[133]^[142] The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks".^[143] A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont.^[144] Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes.^[145] Similar to mycorrhiza, endophytic colonization by fungi may benefit both symbionts; for example, endophytes of grasses impart to their host increased resistance to herbivores and other environmental stresses and receive food and shelter from the plant in return.^[146]

With algae and cyanobacteria

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

Lichens are formed by a symbiotic relationship between algae or cyanobacteria (referred to in lichen terminology as "photobionts") and fungi (mostly various species of ascomycetes and a few basidiomycetes), in which individual photobiont cells are embedded in a tissue formed by the fungus.^[147] Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession,^[148] and are the dominating life forms in extreme environments, including polar, alpine, and semiarid desert regions.^[149] They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves.^[150] As in mycorrhizas, the photobiont provides sugars and other carbohydrates via photosynthesis, while the fungus provides minerals and water. The functions of both symbiotic organisms are so closely intertwined that they function almost as a single organism; in most cases the resulting organism differs greatly from the individual components. Lichenization is a common mode of nutrition; around 20% of fungi—between 17,500 and 20,000 described species—are lichenized.^[151] Characteristics common to most lichens include obtaining organic carbon by photosynthesis, slow growth, small size, long life, long-lasting (seasonal) vegetative reproductive structures, mineral nutrition obtained largely from airborne sources, and greater tolerance of desiccation than most other photosynthetic organisms in the same habitat.^[152]

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Agaricales as their primary food source, while ambrosia beetles cultivate various species of fungi in the bark of trees that they infest.^[153] Similarly, females of several wood wasp species (genus Sirex) inject their eggs together with spores of the wood-rotting fungus Amylostereum areolatum into the sapwood of pine trees; the growth of the fungus provides ideal nutritional conditions for the development of the wasp larvae.^[154] Termites on the African savannah are also known to cultivate fungi,^[135] and yeasts of the genera Candida and Lachancea inhabit the gut of a wide range of insects, including neuropterans, beetles, and cockroaches; it is not known whether these fungi benefit their hosts.^[155] The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae.^[156]

As pathogens and parasites

The plant pathogen Aecidium magellanicum causes calafate rust, seen here on a Berberis shrub in Chile.

Many fungi are parasites on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus Magnaporthe oryzae,^[157] tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease,^[158] and Cryphonectria parasitica responsible for chestnut blight,^[159] and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus.^[139] Some carnivorous fungi, like Paecilomyces lilacinus, are predators of nematodes, which they capture using an array of specialized structures such as constricting rings or adhesive nets.^[160]

Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergilloses, candidoses, coccidioidomycosis, cryptococcosis, histoplasmosis, mycetomas, and paracoccidioidomycosis. Furthermore, persons with immuno-deficiencies are particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus,^[140]^[161]^[162] Histoplasma,^[163] and Pneumocystis.^[164] Other fungi can attack eyes, nails, hair, and especially skin, the so-called dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot.^[165] Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.^[166]

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes. A rapid defense reaction after pathogen attack is the oxidative burst that includes the production of reactive oxygen species at the site of the attempted invasion. As a pathogen, U. maydis can respond to such an oxidative burst by an oxidative stress response, regulated by the gene yap1. This response protects U. maydis from the host attack, and is necessary for the pathogen’s virulence.^[167] Furthermore, U. maydis has a well-established recombinational DNA repair system that acts during mitosis and meiosis.^[168] Recombinational repair during mitosis and meiosis may assist the pathogen in surviving DNA damage arising from the host plant’s oxidative defensive response to infection.^[169]

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. Infection with C. neoformans is termed cryptococcosis, and usually affects the lungs. In the lungs, C. neoformans are phagocytosed by alveolar macrophages.^[170] However some C. neoformans can survive intracellularly in macrophages.^[170] Intracellular survival appears to be the basis for latency, disseminated disease, and resistance to eradication by antifungal agents. One mechanism by which C. neorformans survives the hostile intracellular environment of the macrophage involves up-regulation of expression of genes involved in responses to oxidative stress.^[170] Another mechanism involves meiosis. The great majority of environmental and clinical isolates of C. neoformans are mating type a. Filaments of mating type a ordinarily have haploid nuclei, but these can undergo diploidization (perhaps by endoduplication or by stimulated nuclear fusion) to form diploid cells termed blastospores. The diploid nuclei of blastospores are able to undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed.^[171] This process is referred to as monokaryotic fruiting. Required for this process is a gene, dmc1, a conserved homologue of genes recA in bacteria and rad51 in eukaryotes, that mediates homologous chromosome pairing during meiosis and repair of DNA double-strand breaks (see Meiosis). Thus C. neoformans can undergo a meiotic process, monokaryotic fruiting, that promotes recombinational repair in the oxidative, DNA damaging environment of the host macrophage, and this repair capability may contribute to its virulence.^[171]^[169]

Human use

Saccharomyces cerevisiae cells shown with DIC microscopy

The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history. Mushroom farming and mushroom gathering are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as ethnomycology. Because of the capacity of this group to produce an enormous range of natural products with antimicrobial or other biological activities, many species have long been used or are being developed for industrial production of antibiotics, vitamins, and anti-cancer and cholesterol-lowering drugs. More recently, methods have been developed for genetic engineering of fungi,^[172] enabling metabolic engineering of fungal species. For example, genetic modification of yeast species^[173]—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms.^[174]

Drugs

Many species produce metabolites that are major sources of pharmacologically active drugs. Particularly important are the antibiotics, including the penicillins, a structurally related group of β-lactam antibiotics that are synthesized from small peptides. Although naturally occurring penicillins such as penicillin G (produced by Penicillium chrysogenum) have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by chemical modification of the natural penicillins. Modern penicillins are semisynthetic compounds, obtained initially from fermentation cultures, but then structurally altered for specific desirable properties.^[175] Other antibiotics produced by fungi include: ciclosporin, commonly used as an immunosuppressant during transplant surgery; and fusidic acid, used to help control infection from methicillin-resistant Staphylococcus aureus bacteria.^[176] Widespread use of these antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, and many others began in the early 20th century and continues to play a major part in anti-bacterial chemotherapy. In nature, antibiotics of fungal or bacterial origin appear to play a dual role: at high concentrations they act as chemical defense against competition with other microorganisms in species-rich environments, such as the rhizosphere, and at low concentrations as quorum-sensing molecules for intra- or interspecies signaling.^[177]

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections,^[178] and statins (HMG-CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from Penicillium citrinum and lovastatin from Aspergillus terreus and the oyster mushroom.^[179]

Cultured foods

Baker's yeast or Saccharomyces cerevisiae, a single-celled fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings.^[180] Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation.^[181] Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso,^[182] while Rhizopus species are used for making tempeh.^[183] Several of these fungi are domesticated species that were bred or selected according to their capacity to ferment food without producing harmful mycotoxins (see below), which are produced by very closely related Aspergilli.^[184] Quorn, a meat substitute, is made from Fusarium venenatum.^[185]

Medicinal use

Edible and poisonous species

Amanita phalloides accounts for the majority of fatal mushroom poisonings worldwide.

Edible mushrooms are well-known examples of fungi. Many are commercially raised, but others must be harvested from the wild. Agaricus bisporus, sold as button mushrooms when small or Portobello mushrooms when larger, is a commonly eaten species, used in salads, soups, and many other dishes. Many Asian fungi are commercially grown and have increased in popularity in the West. They are often available fresh in grocery stores and markets, including straw mushrooms (Volvariella volvacea), oyster mushrooms (Pleurotus ostreatus), shiitakes (Lentinula edodes), and enokitake (Flammulina spp.).^[196]

There are many more mushroom species that are harvested from the wild for personal consumption or commercial sale. Milk mushrooms, morels, chanterelles, truffles, black trumpets, and porcini mushrooms (Boletus edulis) (also known as king boletes) demand a high price on the market. They are often used in gourmet dishes.^[197]

Certain types of cheeses require inoculation of milk curds with fungal species that impart a unique flavor and texture to the cheese. Examples include the blue color in cheeses such as Stilton or Roquefort, which are made by inoculation with Penicillium roqueforti.^[198] Molds used in cheese production are non-toxic and are thus safe for human consumption; however, mycotoxins (e.g., aflatoxins, roquefortine C, patulin, or others) may accumulate because of growth of other fungi during cheese ripening or storage.^[199]

Stilton cheese veined with Penicillium roqueforti

Many mushroom species are poisonous to humans, with toxicities ranging from slight digestive problems or allergic reactions as well as hallucinations to severe organ failures and death. Genera with mushrooms containing deadly toxins include Conocybe, Galerina, Lepiota, and most infamously, Amanita.^[200] The latter genus includes the destroying angel (A. virosa) and the death cap (A. phalloides), the most common cause of deadly mushroom poisoning.^[201] The false morel (Gyromitra esculenta) is occasionally considered a delicacy when cooked, yet can be highly toxic when eaten raw.^[202] Tricholoma equestre was considered edible until being implicated in serious poisonings causing rhabdomyolysis.^[203] Fly agaric mushrooms (Amanita muscaria) also cause occasional non-fatal poisonings, mostly as a result of ingestion for use as a recreational drug for its hallucinogenic properties. Historically, fly agaric was used by different peoples in Europe and Asia and its present usage for religious or shamanic purposes is reported from some ethnic groups such as the Koryak people of north-eastern Siberia.^[204]

As it is difficult to accurately identify a safe mushroom without proper training and knowledge, it is often advised to assume that a wild mushroom is poisonous and not to consume it.^[205]^[206]

Pest control

Grasshoppers killed by Beauveria bassiana

In agriculture, fungi may be useful if they actively compete for nutrients and space with pathogenic microorganisms such as bacteria or other fungi via the competitive exclusion principle,^[207] or if they are parasites of these pathogens. For example, certain species may be used to eliminate or suppress the growth of harmful plant pathogens, such as insects, mites, weeds, nematodes and other fungi that cause diseases of important crop plants.^[208] This has generated strong interest in practical applications that use these fungi in the biological control of these agricultural pests. Entomopathogenic fungi can be used as biopesticides, as they actively kill insects.^[209] Examples that have been used as biological insecticides are Beauveria bassiana, Metarhizium spp, Hirsutella spp, Paecilomyces (Isaria) spp, and Lecanicillium lecanii.^[210]^[211] Endophytic fungi of grasses of the genus Neotyphodium, such as N. coenophialum, produce alkaloids that are toxic to a range of invertebrate and vertebrate herbivores. These alkaloids protect grass plants from herbivory, but several endophyte alkaloids can poison grazing animals, such as cattle and sheep.^[212] Infecting cultivars of pasture or forage grasses with Neotyphodium endophytes is one approach being used in grass breeding programs; the fungal strains are selected for producing only alkaloids that increase resistance to herbivores such as insects, while being non-toxic to livestock.^[213]

Bioremediation

Model organisms

Several pivotal discoveries in biology were made by researchers using fungi as model organisms, that is, fungi that grow and sexually reproduce rapidly in the laboratory. For example, the one gene-one enzyme hypothesis was formulated by scientists who used the bread mold Neurospora crassa to test their biochemical theories.^[218] Other important model fungi are Aspergillus nidulans and the yeasts, Saccaromyces cerevisiae and Schizosaccharomyces pombe, each of which has a long history of use to investigate issues in eukaryotic cell biology and genetics, such as cell cycle regulation, chromatin structure, and gene regulation. Other fungal models have more recently emerged that each address specific biological questions relevant to medicine, plant pathology, and industrial uses; examples include Candida albicans, a dimorphic, opportunistic human pathogen,^[219] Magnaporthe grisea, a plant pathogen,^[220] and Pichia pastoris, a yeast widely used for eukaryotic protein expression.^[221]

Others

Fungi are used extensively to produce industrial chemicals like citric, gluconic, lactic, and malic acids,^[222] and industrial enzymes, such as lipases used in biological detergents,^[223] cellulases used in making cellulosic ethanol^[224] and stonewashed jeans,^[225] and amylases,^[226] invertases, proteases and xylanases.^[227] Several species, most notably Psilocybin mushrooms (colloquially known as magic mushrooms), are ingested for their psychedelic properties, both recreationally and religiously.

Mycotoxins

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

Many fungi produce biologically active compounds, several of which are toxic to animals or plants and are therefore called mycotoxins. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal amatoxins in some Amanita mushrooms, and ergot alkaloids, which have a long history of causing serious epidemics of ergotism (St Anthony's Fire) in people consuming rye or related cereals contaminated with sclerotia of the ergot fungus, Claviceps purpurea.^[228] Other notable mycotoxins include the aflatoxins, which are insidious liver toxins and highly carcinogenic metabolites produced by certain Aspergillus species often growing in or on grains and nuts consumed by humans, ochratoxins, patulin, and trichothecenes (e.g., T-2 mycotoxin) and fumonisins, which have significant impact on human food supplies or animal livestock.^[229]

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi.^[28] Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory).^[230]^[231]

Mycology

Mycology is the branch of biology concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and psychotropic substances consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi.^[232]

In 1729, Pier A. Micheli first published descriptions of fungi.

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus).^[233] Ancient peoples have used fungi as food sources–often unknowingly–for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.^[234]

History

Mycology is a relatively new science that became systematic after the development of the microscope in the 16th century. Although fungal spores were first observed by Giambattista della Porta in 1588, the seminal work in the development of mycology is considered to be the publication of Pier Antonio Micheli's 1729 work Nova plantarum genera.^[235] Micheli not only observed spores, but showed that under the proper conditions, they could be induced into growing into the same species of fungi from which they originated.^[236] Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christian Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill so as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and various microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. The 20th century has seen a modernization of mycology that has come from advances in biochemistry, genetics, molecular biology, and biotechnology. The use of DNA sequencing technologies and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.^[237]

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^ El Dine RS, El Halawany AM, Ma CM, Hattori M. (2008). "Anti-HIV-1 protease activity of lanostane triterpenes from the vietnamese mushroom Ganoderma colossum". Journal of Natural Products 71 (6): 1022–1026. doi:10.1021/np8001139. PMID 18547117.
^ Yuen JW, Gohel MD. (2005). "Anticancer effects of Ganoderma lucidum: a review of scientific evidence". Nutrition and Cancer 53 (1): 11–17. doi:10.1207/s15327914nc5301_2. PMID 16351502.
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^ Halpern GM, Miller A. (2002). Medicinal Mushrooms: Ancient Remedies for Modern Ailments. New York, New York: M. Evans and Co. p. 116. ISBN 0-87131-981-0.
^ Fisher M, Yang LX. (2002). "Anticancer effects and mechanisms of polysaccharide-K (PSK): implications of cancer immunotherapy". Anticancer Research 22 (3): 1737–1754. PMID 12168863.
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^ Hall, pp. 13–26.
^ Kinsella JE, Hwang DH. (1976). "Enzymes of Penicillium roqueforti involved in the biosynthesis of cheese flavor". CRC Critical Reviews in Food Science and Nutrition 8 (2): 191–228. doi:10.1080/10408397609527222. PMID 21770.
^ Erdogan A, Gurses M, Sert S. (2004). "Isolation of moulds capable of producing mycotoxins from blue mouldy Tulum cheeses produced in Turkey". International Journal of Food Microbiology 85 (1–2): 83–85. doi:10.1016/S0168-1605(02)00485-3. PMID 12810273.
^ Orr DB, Orr RT. (1979). Mushrooms of Western North America. Berkeley, California: University of California Press. p. 17. ISBN 0-520-03656-5.
^ Vetter J. (1998). "Toxins of Amanita phalloides". Toxicon 36 (1): 13–24. doi:10.1016/S0041-0101(97)00074-3. PMID 9604278.
^ Leathem AM, Dorran TJ. (2007). "Poisoning due to raw Gyromitra esculenta (false morels) west of the Rockies". Canadian Journal of Emergency Medicine 9 (2): 127–130. PMID 17391587.
^ Karlson-Stiber C, Persson H. (2003). "Cytotoxic fungi—an overview". Toxicon 42 (4): 339–349. doi:10.1016/S0041-0101(03)00238-1. PMID 14505933.
^ Michelot D, Melendez-Howell LM. (2003). "Amanita muscaria: chemistry, biology, toxicology, and ethnomycology". Mycological Research 107 (Pt 2): 131–146. doi:10.1017/S0953756203007305. PMID 12747324.
^ Hall, p. 7.
^ Ammirati JF, McKenny M, Stuntz DE. (1987). The New Savory Wild Mushroom. Seattle, Washington: University of Washington Press. pp. xii–xiii. ISBN 0-295-96480-4.
^ López-Gómez J, Molina-Meyer M. (2006). "The competitive exclusion principle versus biodiversity through competitive segregation and further adaptation to spatial heterogeneities". Theoretical Population Biology 69 (1): 94–109. doi:10.1016/j.tpb.2005.08.004. PMID 16223517.
^ Becker H. (1998). "Setting the Stage To Screen Biocontrol Fungi". United States Department of Agriculture, Agricultural Research Service. Retrieved 2009-02-23.
^ Keiller TS. "Whey-based fungal microfactory technology for enhanced biological pest management using fungi" (PDF). UVM Innovations. Retrieved 2011-10-29.
^ Deshpande MV. (1999). "Mycopesticide production by fermentation: potential and challenges". Critical Reviews in Microbiology 25 (3): 229–243. doi:10.1080/10408419991299220. PMID 10524330.
^ Thomas MB, Read AF. (2007). "Can fungal biopesticides control malaria?". Nature Reviews Microbiology 5 (5): 377–383. doi:10.1038/nrmicro1638. PMID 17426726.
^ Bush LP, Wilkinson HH, Schardl CL. (1997). "Bioprotective alkaloids of grass-fungal endophyte symbioses". Plant Physiology 114 (1): 1–7. PMC 158272. PMID 12223685.
^ Bouton JH, Latch GCM, Hill NS, Hoveland CS, McCannc MA, Watson RH, Parish JA, Hawkins LL, Thompson FN. (2002). "Use of nonergot alkaloid-producing endophytes for alleviating tall fescue toxicosis in sheep". Agronomy Journal 94: 567–574. review
^ Christian V, Shrivastava R, Shukla D, Modi HA, Vyas BR. (2005). "Degradation of xenobiotic compounds by lignin-degrading white-rot fungi: enzymology and mechanisms involved" (PDF). Indian Journal of Experimental Biology 43 (4): 301–312. PMID 15875713. ^{[dead link]}
^ "Fungi to fight 'toxic war zones'". BBC News. May 5, 2008.
^ Fomina M, Charnock JM, Hillier S, Alvarez R, Gadd GM. (2007). "Fungal transformations of uranium oxides". Environmental Microbiology 9 (7): 1696–1710. doi:10.1111/j.1462-2920.2007.01288.x. PMID 17564604.
^ Fomina M, Charnock JM, Hillier S, Alvarez R, Livens F, Gadd GM. (2008). "Role of fungi in the biogeochemical fate of depleted uranium". Current Biology 18 (9): R375–R377. doi:10.1016/j.cub.2008.03.011. PMID 18460315.
^ Beadle GW, Tatum EL. (1941). "Genetic control of biochemical reactions in Neurospora". Proceedings of the National Academy of Sciences of the USA 27 (11): 499–506. Bibcode:1941PNAS...27..499B. doi:10.1073/pnas.27.11.499. PMC 1078370. PMID 16588492.
^ Datta A, Ganesan K, Natarajan K. (1989). "Current trends in Candida albicans research". Advances in Microbial Physiology. Advances in Microbial Physiology 30: 53–88. doi:10.1016/S0065-2911(08)60110-1. ISBN 978-0-12-027730-8. PMID 2700541.
^ Dean RA. (2005). "The genome sequence of the rice blast fungus Magnaporthe grisea". Nature 434 (7036): 980–986. Bibcode:2005Natur.434..980D. doi:10.1038/nature03449. PMID 15846337.
^ Daly R, Hearn MT. (2005). "Expression of heterologous proteins in Pichia pastoris: a useful experimental tool in protein engineering and production". Journal of Molecular Recognition 18 (2): 119–138. doi:10.1002/jmr.687. PMID 15565717.
^ Schlegel HG. (1993). General Microbiology. Cambridge, UK: Cambridge University Press. p. 360. ISBN 0-521-43980-9.
^ Joseph B, Ramteke PW, Thomas G. (2008). "Cold active microbial lipases: some hot issues and recent developments". Biotechnology Advances 26 (5): 457–470. doi:10.1016/j.biotechadv.2008.05.003. PMID 18571355.
^ Kumar R, Singh S, Singh OV. (2008). "Bioconversion of lignocellulosic biomass: biochemical and molecular perspectives". Journal of Industrial Microbiology and Biotechnology 35 (5): 377–391. doi:10.1007/s10295-008-0327-8. PMID 18338189.
^ "Trichoderma spp., including T. harzianum, T. viride, T. koningii, T. hamatum and other spp. Deuteromycetes, Moniliales (asexual classification system)". Biological Control: A Guide to Natural Enemies in North America. Retrieved 2007-07-10.
^ Olempska-Beer ZS, Merker RI, Ditto MD, DiNovi MJ. (2006). "Food-processing enzymes from recombinant microorganisms—a review". Regulatory Toxicology and Pharmacology 45 (2): 144–158. doi:10.1016/j.yrtph.2006.05.001. PMID 16769167.
^ Polizeli ML, Rizzatti AC, Monti R, Terenzi HF, Jorge JA, Amorim DS. (2005). "Xylanases from fungi: properties and industrial applications". Applied Microbiology and Biotechnology 67 (5): 577–591. doi:10.1007/s00253-005-1904-7. PMID 15944805.
^ Schardl CL, Panaccione DG, Tudzynski P. (2006). "Ergot alkaloids—biology and molecular biology". The Alkaloids. Chemistry and Biology. The Alkaloids: Chemistry and Biology 63: 45–86. doi:10.1016/S1099-4831(06)63002-2. ISBN 978-0-12-469563-4. PMID 17133714.
^ van Egmond HP, Schothorst RC, Jonker MA. (2007). "Regulations relating to mycotoxins in food: perspectives in a global and European context". Analytical and Bioanalytical Chemistry 389 (1): 147–157. doi:10.1007/s00216-007-1317-9. PMID 17508207.
^ Demain AL, Fang A. (2000). "The natural functions of secondary metabolites". Advances in Biochemical Engineering/Biotechnology. Advances in Biochemical Engineering/Biotechnology 69: 1–39. doi:10.1007/3-540-44964-7_1. ISBN 978-3-540-67793-2. PMID 11036689.
^ Rohlfs M, Albert M, Keller NP, Kempken F. (2007). "Secondary chemicals protect mould from fungivory". Biology Letters 3 (5): 523–525. doi:10.1098/rsbl.2007.0338. PMC 2391202. PMID 17686752.
^ According to one 2001 estimate, some 10,000 fungal diseases are known. Struck C. (2006). "Infection strategies of plant parasitic fungi". In Cooke BM, Jones DG, Kaye B. The Epidemiology of Plant Diseases. Berlin, Germany: Springer. p. 117. ISBN 1-4020-4580-8.
^ Peintner U, Pöder R, Pümpel T. (1998). "The Iceman's fungi". Mycological Research 102 (10): 1153–1162. doi:10.1017/S0953756298006546.
^ Ainsworth, p. 1.
^ Alexopoulos et al., pp. 1–2.
^ Ainsworth, p. 18.
^ Hawksworth DL. (2006). "Pandora's Mycological Box: Molecular sequences vs. morphology in understanding fungal relationships and biodiversity". Revista Iberoamericana de Micologia 23 (3): 127–133. doi:10.1016/S1130-1406(06)70031-6. PMID 17196017.

Cited literature

Ainsworth GC. (1976). Introduction to the History of Mycology. Cambridge, UK: Cambridge University Press. ISBN 0-521-11295-8.
Alexopoulos CJ, Mims CW, Blackwell M. (1996). Introductory Mycology. John Wiley and Sons. ISBN 0-471-52229-5.
Chandler PJ. (2010). A Dipterist's Handbook (2nd Edition). The Amateur Entomologists' Society. pp. 1–525.
Deacon J. (2005). Fungal Biology. Cambridge, Massachusetts: Blackwell Publishers. ISBN 1-4051-3066-0.
Hall IR. (2003). Edible and Poisonous Mushrooms of the World. Portland, Oregon: Timber Press. ISBN 0-88192-586-1.
Hanson JR. (2008). The Chemistry of Fungi. Royal Society Of Chemistry. ISBN 0-85404-136-2.
Jennings DH, Lysek G. (1996). Fungal Biology: Understanding the Fungal Lifestyle. Guildford, UK: Bios Scientific Publishers Ltd. ISBN 978-1-85996-150-6.
Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi. 10th ed. Wallingford, UK: CAB International. ISBN 0-85199-826-7.
Taylor EL, Taylor TN. (1993). The Biology and Evolution of Fossil Plants. Englewood Cliffs, New Jersey: Prentice Hall. ISBN 0-13-651589-4.

External links

Look up fungus in Wiktionary, the free dictionary.

Find more about Fungus at Wikipedia's sister projects
	Definitions and translations from Wiktionary
	Media from Commons
	Learning resources from Wikiversity
	Quotations from Wikiquote
	Source texts from Wikisource
	Textbooks from Wikibooks
	Species directory on Fungi from Wikispecies

Tree of Life web project: Fungi
Fungus at the Encyclopedia of Life
Mushroom Observer (mushroomobserver.org), a collaborative fungus recording and identification project

UpToDate Contents

全文を閲覧するには購読必要です。 To read the full text you will need to subscribe.

1. ヒトの疾患における真菌（カビ）の役割 the role of fungi molds in human disease
2. 難治性肺炎 nonresolving pneumonia
3. 黒色真菌による中枢神経系の感染症（脳黒色菌糸症） central nervous system infections due to dematiaceous fungi cerebral phaeohyphomycosis
4. 抗真菌剤感受性試験 antifungal susceptibility testing
5. 頭部白癬 tinea capitis

English Journal

Improved endoglucanase production and mycelial biomass of some ericoid fungi.

Adeoyo OR1, Pletschke BI1, Dames JF2.
AMB Express.AMB Express.2017 Dec;7(1):15. doi: 10.1186/s13568-016-0312-y. Epub 2017 Jan 3.
Fungal species associated with ericaceous plant roots produce a number of enzymes and other bio-active metabolites in order to enhance survival of their host plants in natural environments. This study focussed on endoglucanase production from root associated ericoid mycorrhizal and dark septate endo
PMID 28050856

Selection of reference genes from Shiraia bambusicola for RT-qPCR analysis under different culturing conditions.

Zhang C1, Li T1, Hou CL2, Shen XY3.
AMB Express.AMB Express.2017 Dec;7(1):14. doi: 10.1186/s13568-016-0314-9. Epub 2017 Jan 3.
Stable reference genes are necessary to analyse quantitative real-time reverse transcription PCR (qRT-PCR) data and determine the reliability of the final results. For further studies of the valuable fungus Shiraia bambusicola, the identification of suitable reference genes has become increasingly u
PMID 28050855

Metagenomic insights into the rumen microbial fibrolytic enzymes in Indian crossbred cattle fed finger millet straw.

Jose VL1,2, Appoothy T3, More RP4, Arun AS5.
AMB Express.AMB Express.2017 Dec;7(1):13. doi: 10.1186/s13568-016-0310-0. Epub 2017 Jan 3.
The rumen is a unique natural habitat, exhibiting an unparalleled genetic resource of fibrolytic enzymes of microbial origin that degrade plant polysaccharides. The objectives of this study were to identify the principal plant cell wall-degrading enzymes and the taxonomic profile of rumen microbial
PMID 28050853

Japanese Journal

Cytochalasin H, an Active Anti-Angiogenic Constituent of the Ethanol Extract of Gleditsia sinensis Thorns

Lee Jun,Yi Jin-Mu,Kim Haejin,Lee You Jin,Park Jong-Shik,Bang Ok-Sun,Kim No Soo
Biological and Pharmaceutical Bulletin, 2014
… By virtue of in vitro activity-guided fractionation using human umbilical vein endothelial cells (HUVEC) primary endothelial cells, chromatographic separation, and NMR spectral analyses, we isolated and identified the potent active constituent, cytochalasin H, a biologically active secondary metabolite of fungi. … This unexpected active constituent may have originated from the endophytic fungi, Chaetomium globosum, which naturally populate G. …
NAID 130003382111

De Novo Design of Non-coordinating Indolones as Potential Inhibitors for Lanosterol 14-α-Demethylase (CYP51)

González-Chávez Rodolfo,Martínez Roberto,Torre-Bouscoulet María Eugenia,Gallo Marco,González-Chávez Marco Martín
Chemical and Pharmaceutical Bulletin 62(1), 16-24, 2014
… The antifungal activities of these derivatives against eight human pathogenic filamentous fungi and yeast strains were evaluated in vitro by measuring the minimal inhibitory concentrations. …
NAID 130003381800

糸状菌の極性生長 : 細胞骨格と形質膜の相互作用による極性の制御

竹下典男
化学と生物 : 日本農芸化学会会誌 : 生命・食糧・環境 51(12), 809-820, 2013-12
NAID 40019902106

Related Pictures

★リンクテーブル★

リンク元	「真菌」「microbial」「菌類」「菌」「真菌類」
拡張検索	「fungivorous」「minimal fungicidal concentration」

「真菌」

Library resources
About Fungus
	Online books; Resources in your library; Resources in other libraries;

Fungi; Temporal range: Early Devonian–Recent (but see text) PreЄ Є O S D C P T J K Pg N
	Clockwise from top left:; Amanita muscaria, a basidiomycete;; Sarcoscypha coccinea, an ascomycete;; bread covered in mold;; a chytrid;; an Aspergillus conidiophore.
Scientific classification
Domain:	Eukaryota
(unranked):	Opisthokonta
Kingdom:	Fungi; (L., 1753) R.T. Moore, 1980
Subkingdoms/Phyla/Subphyla
	Blastocladiomycota Chytridiomycota Glomeromycota Microsporidia Neocallimastigomycota Dikarya (inc. Deuteromycota) Ascomycota Pezizomycotina Saccharomycotina Taphrinomycotina Basidiomycota Agaricomycotina Pucciniomycotina Ustilaginomycotina Subphyla incertae sedis Entomophthoromycotina Kickxellomycotina Mucoromycotina Zoopagomycotina

　　[★]

英: fungus, (pl.)fungi
同: かび、菌類 Mycota
関: 微生物学、真菌症、抗真菌薬

細菌、真菌、藻類、原虫

いずれも単細胞生物
細菌は核を持たないが、それ以外は核を持つ
細胞壁の多糖：細菌はペプチドグリカンなど。真菌はβグルカン、キチン。藻類はセルロース。原虫は細胞壁を持たない
栄養獲得様式：藻類は光合成独立栄養生性。

大きさ

数μm。観察に油浸レンズは不要

細胞膜

リン脂質＋グリセリド＋ステロール(6%。主にエルゴステロール)
機能：細胞内への物質輸送、細胞内物質の排出、細胞内浸透圧調節
細胞壁の合成酵素(β-1,3-グルカン合成酵素,キチン合成酵素)・分解酵素(キチナーゼ、β-グルカナーゼ)が存在

細胞壁

線維状多糖＋糖タンパク質
真菌に特有の成分(β-グルカン、マンナン、ガラクトマンナン)から構成される

線維状多糖

細胞壁の骨格となる
キチンとβ-グルカンからなる。
特殊な多糖にキトサンがある

キチン

N-アセチルグルコサミンのホモ重合体
β(1→4)結合
リゾチームで分解される
糸状菌では細胞壁成分にキチンが多い

β-グルカン

グルコースのホモ重合体
β(1→3)結合、β(1→6)結合
酵母では細胞壁成分にβ-1,3-グルカンが多い

キトサン

グルコサミンのホモ重合体
接合菌はキチンの代わりにキトサンを多く含む一方、β-グルカンをほとんど含まない。

糖タンパク質

線維状多糖の間を埋めている糖タンパク質
アスペルギルス属：ガラクトマンナン
カンジダ属　　　：マンナン。α(1→6)結合からなる主鎖。α(1→2)結合とα(1→3)結合からなるD-マンノースのホモ重合体

形態による分類

糸状菌 filamentous fungi 菌糸状真菌 mycelial fungi
酵母 yeast 酵母状真菌 yeastlike fungi
二形性真菌 dimorphic fungi

Coccidioides immitis

Histoplasma capsulatum

Sporothrix schenckii

Candida albicans

培養と感染組織における形態

カンジダ属 Candida albicans　：　培養(球形)　組織内(仮性菌糸)
アスペルギルス属 Aspergillus fumigatus　：　培地(胞子産生)　組織内(菌糸. 胞子は末梢に出てこない)
クリプトコッカス属 Cryptococcus neoformans　：　培地(球形)　組織内(球形)

菌糸の構造

無隔菌糸

接合菌などの下等真菌にのみ見られる

有隔菌糸
高等真菌(子嚢菌、担子菌、不完全菌)

菌糸の機能

栄養菌糸 vegetative hypha 基質菌糸 substrate hypha
生殖菌糸 reproductive hypha

胞子を形成

分生子柄：形成される胞子が分生子である場合の生殖菌糸の特別な呼称

生殖方式による分類

有性生殖と無性生殖

完全菌 perfect fungi

有性生殖と無性生殖を行う

有性生殖により形成された胞子：有性胞子 sexual spore
無性生殖により形成された胞子：無性胞子 asexual spore

不完全菌 imperfect fungi

無性生殖のみ行う

有性胞子形成

無性胞子形成

培養

サブロー･グルコース寒天培地 Sabouraud glucose agar

グルコース1-4%, ペプトン1%

種類

真菌の染色法(SMB.358)

細胞壁の多糖を染色：コットンブルー(cotton blue)、グラム染色(全ての真菌はグラム陽性)
真菌細胞壁多糖を特異的に染色：PAS染色、Grocottメテナミン銀染色、ファンギフローラY

参考

真菌の写真 - Fungal Images

http://www.asm.org/Division/c/fungi.htm

病原体としての真菌

分類群

菌名

病名

出芽分裂

菌体

菌糸

胞子

発芽管

寄生性

感染経路

病原体の特徴

病型

検査

治療

Candida albicans

形成

常在菌→各所

日和見感染症

担子菌

Filobasidiella neoformans

クリプトコッカス症

出芽

厚い莢膜

通性細胞内寄生体

鳩の乾燥糞→空気感染→肺(通性細胞内寄生体)

日和見感染菌

肺クリプトコッカス症、
中枢神経クリプトコッカス症、
皮膚クリプトコッカス症、
全身性クリプトコッカス症

墨汁染色

( アムホテリシンB or フルコナゾール )
　 ± フルシトシン

不完全菌

Trichosporon asahii

トリコスポロン症

仮性菌糸

分節分生子

子嚢菌

Aspergillus fumigatus/
'Aspergillus flavus

アスペルギルス症

糸状菌
不完全菌

有隔菌糸

土壌や朽ちた植物、空気中の浮遊胞子

Mucor/'Rhizopus/
Absidia/Rhizomucor

ムコール症

無隔菌糸

血管に親和性

土壌や朽ちた植物、空気中の浮遊胞子
　→　浮遊胞子の吸入

鼻眼脳型ムコール症
侵襲性肺ムコール症
全身播種型ムコール症

β-D-グルカン陰性。
喀痰、血培陰性

アムホテリシンB

子嚢菌

Pneumocystis jirovecii
('Pneumocystis carini)

ニューモシスチス・カリニ感染症

子嚢菌

Histoplasma capsulatum

ヒストプラスマ症

二形性真菌
　37℃酵母型発育
　25℃菌糸形成

空気感染。肺に一次病変を作り様々な臓器に播種

AIDS患者に好発

不完全菌

Coccidioides immitis

コクシジオイデス症

二形性真菌

内生胞子

不完全菌

Sporothrix schenckii

スポロトリクス症

二形性真菌

「microbial」

　　[★]

adj.

微生物の、菌の

関: fungi、fungus、germ、microbe、microorganism

「菌類」

　　[★]

英: fungus、fungi, Mycota
関: カビ、糸状菌、真菌、真菌類、菌

「菌」

　　[★]

英: fungus、fungi、microbial
関: 菌類、真菌、真菌類、微生物

「真菌類」

　　[★]

英: fungus、fungi
関: 菌類、真菌、菌

「fungivorous」

　　[★]

菌食性の

関: mycophagy

「minimal fungicidal concentration」

　　[★] 最小殺真菌濃度 MFC

[Moore1980-1] Moore RT. (1980). "Taxonomic proposals for the classification of marine yeasts and other yeast-like fungi including the smuts". Botanica Marine 23: 361–373.

[2] The classification system presented here is based on the 2007 phylogenetic study by Hibbett et al.

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Fungi Temporal range: Early Devonian–Recent (but see text) PreЄ Є O S D C P T J K Pg N

Clockwise from top left: Amanita muscaria, a basidiomycete; Sarcoscypha coccinea, an ascomycete; bread covered in mold; a chytrid; an Aspergillus conidiophore.
Scientific classification
Domain:	Eukaryota
(unranked):	Opisthokonta
Kingdom:	Fungi (L., 1753) R.T. Moore, 1980^[1]
Subkingdoms/Phyla/Subphyla^[2]
Blastocladiomycota Chytridiomycota Glomeromycota Microsporidia Neocallimastigomycota Dikarya (inc. Deuteromycota) Ascomycota Pezizomycotina Saccharomycotina Taphrinomycotina Basidiomycota Agaricomycotina Pucciniomycotina Ustilaginomycotina Subphyla incertae sedis Entomophthoromycotina Kickxellomycotina Mucoromycotina Zoopagomycotina

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Contents

Etymology

Characteristics

Diversity

Morphology

Microscopic structures

Macroscopic structures

Growth and physiology

Reproduction

Asexual reproduction

Sexual reproduction

Spore dispersal

Other sexual processes

Evolution

Taxonomy

Taxonomic groups

Fungus-like organisms

Ecology

Symbiosis

With plants

With algae and cyanobacteria

With insects

As pathogens and parasites

Pathogenic mechanisms

Human use

Drugs

Cultured foods

Medicinal use

Edible and poisonous species

Pest control

Bioremediation

Model organisms

Others

Mycotoxins

Mycology

History

See also

References

Cited literature

External links

UpToDate Contents

English Journal

Japanese Journal

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★リンクテーブル★

「真菌」

細菌、真菌、藻類、原虫

大きさ

細胞膜

細胞壁

線維状多糖

キチン

β-グルカン

キトサン

糖タンパク質

形態による分類

培養と感染組織における形態

菌糸の構造

菌糸の機能

生殖方式による分類

有性生殖と無性生殖

有性胞子形成

無性胞子形成

培養

種類

真菌の染色法(SMB.358)

参考

病原体としての真菌

「microbial」

「菌類」

「菌」