thumb|upright=1.35|[[Pholiota squarrosa growing at the base of a tree]]

A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or another food source. A toadstool generally refers to a poisonous mushroom.

The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus; hence, the word "mushroom" is most often applied to those fungi (Basidiomycota, Agaricomycetes) that have a stem (stipe), a cap (pileus), and gills (lamellae, sing. lamella) on the underside of the cap. "Mushroom" also describes a variety of other gilled fungi, with or without stems; therefore the term is used to describe the fleshy fruiting bodies of some Ascomycota. The gills produce microscopic spores which help the fungus spread across the ground or its occupant surface.

Forms deviating from the standard morphology usually have more specific names, such as "bolete", "truffle", "puffball", "stinkhorn", and "morel", and gilled mushrooms themselves are often called "agarics" in reference to their similarity to Agaricus or their order Agaricales.

Mushrooms grow from underground mycelium and can expand rapidly under favorable conditions. They vary widely in use: some are edible and nutritious, while others are toxic or psychoactive; many have roles in folk medicine, ecology, and industry.

Etymology

thumb|upright=0.85|[[Amanita muscaria, the most easily recognised "toadstool", is frequently depicted in fairy stories and on greeting cards. It is often associated with gnomes.]]

The terms "mushroom" and "toadstool" go back centuries and were never precisely defined, nor was there consensus on application. During the 15th and 16th centuries, the terms mushrom, mushrum, muscheron, mousheroms, mussheron, or musserouns were used. The word toadstool appeared first in 14th-century England as a reference for a "stool" for toads, possibly implying an inedible poisonous fungus, an association it carries in modern terminology.

Identification

Identifying what is and is not a mushroom requires a basic understanding of their macroscopic structure. Most are basidiomycetes and gilled. Their spores, called basidiospores, are produced on the gills and fall in a fine rain of powder from under the caps as a result. At the microscopic level, the basidiospores are shot off basidia and then fall between the gills in the dead air space. As a result, for most mushrooms, if the cap is cut off and placed gill-side-down overnight, a powdery impression reflecting the shape of the gills (or pores, or spines, etc.) is formed (when the fruit body is sporulating). The color of the powdery print, called a spore print, is useful in both classifying and identifying mushrooms. Spore print colors include white (most common), brown, black, purple-brown, pink, yellow, and creamy, but almost never blue, green, or red.

In general, identification to genus can often be accomplished in the field using a local field guide. Identification to species, however, requires more effort. A mushroom develops from a button stage into a mature structure, and only the latter can provide certain characteristics needed for the identification of the species. However, over-mature specimens lose features and cease producing spores. Many novices have mistaken humid water marks on paper for white spore prints, or discolored paper from oozing liquids on lamella edges for colored spored prints.

Classification

thumb|A mushroom (probably [[Russula brevipes) parasitized by Hypomyces lactifluorum resulting in a "lobster mushroom"]]

Typical mushrooms are the fruit bodies of members of the order Agaricales, whose type genus is Agaricus and type species is the field mushroom, Agaricus campestris. However in modern molecularly defined classifications, not all members of the order Agaricales produce mushroom fruit bodies, and many other gilled fungi, collectively called mushrooms, occur in other orders of the class Agaricomycetes. For example, chanterelles are in the Cantharellales, false chanterelles such as Gomphus are in the Gomphales, milk-cap mushrooms (Lactarius, Lactifluus) and russulas (Russula), as well as Lentinellus, are in the Russulales, while the tough, leathery genera Lentinus and Panus are among the Polyporales, but Neolentinus is in the Gloeophyllales, and the little pin-mushroom genus, Rickenella, along with similar genera, are in the Hymenochaetales.

Within the main body of mushrooms, in the Agaricales, are common fungi like the common fairy-ring mushroom, shiitake, enoki, oyster mushrooms, fly agarics and other Amanitas, magic mushrooms like species of Psilocybe, paddy straw mushrooms, shaggy manes, etc.

An atypical mushroom is the lobster mushroom, which is a fruit body of a Russula or Lactarius mushroom that has been deformed by the parasitic fungus Hypomyces lactifluorum. This gives the affected mushroom an unusual shape and red color that resembles that of a boiled lobster.

The stalk (also called the stipe, or stem) may be central and support the cap in the middle, or it may be off-center or lateral, as in species of Pleurotus and Panus. In other mushrooms, a stalk may be absent, as in the polypores that form shelf-like brackets. Puffballs lack a stalk, but may have a supporting base. Other mushrooms including truffles, jellies, earthstars, and bird's nests usually do not have stalks, and a specialized mycological vocabulary exists to describe their parts.

The way the gills attach to the top of the stalk is an important feature of mushroom morphology. Mushrooms in the genera Agaricus, Amanita, Lepiota and Pluteus, among others, have free gills that do not extend to the top of the stalk. Others have decurrent gills that extend down the stalk, as in the genera Omphalotus and Pleurotus. There are a great number of variations between the extremes of free and decurrent, collectively called attached gills. Finer distinctions are often made to distinguish the types of attached gills: adnate gills, which adjoin squarely to the stalk; notched gills, which are notched where they join the top of the stalk; adnexed gills, which curve upward to meet the stalk, and so on. These distinctions between attached gills are sometimes difficult to interpret, since gill attachment may change as the mushroom matures, or with different environmental conditions.

Microscopic features

A hymenium is a layer of microscopic spore-bearing cells that covers the surface of gills. In the nongilled mushrooms, the hymenium lines the inner surfaces of the tubes of boletes and polypores, or covers the teeth of spine fungi and the branches of corals. In the Ascomycota, spores develop within microscopic elongated, sac-like cells called asci, which typically contain eight spores in each ascus. The Discomycetes, which contain the cup, sponge, brain, and some club-like fungi, develop an exposed layer of asci, as on the inner surfaces of cup fungi or within the pits of morels. The Pyrenomycetes, tiny dark-colored fungi that live on a wide range of substrates including soil, dung, leaf litter, and decaying wood, as well as other fungi, produce minute, flask-shaped structures called perithecia, within which the asci develop.

In the basidiomycetes, usually four spores develop on the tips of thin projections called sterigmata, which extend from club-shaped cells called a basidia. The fertile portion of the Gasteromycetes, called a gleba, may become powdery as in the puffballs or slimy as in the stinkhorns. Interspersed among the asci are threadlike sterile cells called paraphyses. Similar structures called cystidia often occur within the hymenium of the Basidiomycota. Many types of cystidia exist, and assessing their presence, shape, and size is often used to verify the identification of a mushroom. Glucans have potential roles in reserving energy, providing structure, cell-to-cell signaling, and cellular protection.

The cultivated mushroom, as well as the common field mushroom, initially form a minute fruiting body, referred to as the pin stage because of their small size. Slightly expanded, they are called buttons, once again because of the relative size and shape. Once such stages are formed, the mushroom can rapidly pull in water from its mycelium and expand, mainly by inflating preformed cells that took several days to form in the primordia.

Similarly, there are other mushrooms, like Parasola plicatilis (formerly Coprinus plicatlis), that grow rapidly overnight and may disappear by late afternoon on a hot day after rainfall.

Not all mushrooms expand overnight; some grow very slowly and add tissue to their fruiting bodies by growing from the edges of the colony or by inserting hyphae. For example, Pleurotus nebrodensis grows slowly, and because of this combined with human collection, it is now critically endangered. Most of the fungus is underground and in decaying wood or dying tree roots in the form of white mycelia combined with black shoelace-like rhizomorphs that bridge colonized separated woody substrates. When exposed to ultraviolet (UV) light, even after harvesting, ergosterol in mushrooms is converted to vitamin D<sub>2</sub>, In a comprehensive safety assessment of producing vitamin D in fresh mushrooms, researchers showed that artificial UV light technologies were equally effective for vitamin D production as in mushrooms exposed to natural sunlight, and that UV light has a long record of safe use for production of vitamin D in food.

Human use

Edible mushrooms

thumb|left|[[Agaricus bisporus, one of the most widely cultivated and consumed mushrooms]]

thumb|left| [[Pleurotus eryngii var. ferulae|Ferula mushroom in Bingöl, Turkey. This is an edible type of mushroom.]]

Mushrooms are used extensively in cooking, in many cuisines (notably Chinese, Korean, European, and Japanese). Humans have valued them as food since antiquity.

Most mushrooms sold in supermarkets have been commercially grown on mushroom farms. The most common of these, Agaricus bisporus, is considered safe for most people to eat because it is grown in controlled, sterilized environments. Several varieties of A.&nbsp;bisporus are grown commercially, including whites, crimini, and portobello. Other cultivated species available at many grocers include Hericium erinaceus, shiitake, maitake (hen-of-the-woods), Pleurotus, and enoki. In recent years, increasing affluence in developing countries has led to a considerable growth in interest in mushroom cultivation, which is now seen as a potentially important economic activity for small farmers. The country produces about half of all cultivated mushrooms, and around of mushrooms are consumed per person per year by 1.4&nbsp;billion people. In 2014, Poland was the world's largest mushroom exporter, reporting an estimated annually.

Separating edible from poisonous species requires meticulous attention to detail; there is no single trait by which all toxic mushrooms can be identified, nor one by which all edible mushrooms can be identified. People who collect mushrooms for consumption are known as mycophagists, Even the cultivated A.&nbsp;bisporus contains small amounts of hydrazines, the most abundant of which is agaritine (a mycotoxin and carcinogen). However, the hydrazines are destroyed by moderate heat when cooking.

In Jainism, the consumption of mushrooms and other edible fungi is strictly prohibited. This dietary restriction is fundamentally rooted in the religion's core ethical principle of nonviolence (). Classical Jain taxonomy classifies mushrooms as , meaning they are considered to be bodies that harbor an infinite number of microscopic souls or lifeforms (). Furthermore, because mushrooms frequently grow in dark environments or on decaying organic matter, traditional Jain biology views them as continuous breeding grounds for mobile microorganisms (). Consequently, Jain dietary laws mandate the absolute avoidance of mushrooms to prevent the mass destruction of these microscopic organisms.

Toxic mushrooms

thumb|Young [[Amanita phalloides "death cap" mushrooms, with a matchbox for size comparison]]

Many mushroom species produce secondary metabolites that can be toxic, mind-altering, antibiotic, antiviral, or bioluminescent. Although there are only a small number of deadly species, several others can cause particularly severe and unpleasant symptoms. Toxicity likely plays a role in protecting the function of the basidiocarp: the mycelium has expended considerable energy and protoplasmic material to develop a structure to efficiently distribute its spores. One defense against consumption and premature destruction is the evolution of chemicals that render the mushroom inedible, either causing the consumer to vomit the meal (see emetics), or to learn to avoid consumption altogether. In addition, due to the propensity of mushrooms to absorb heavy metals, including those that are radioactive, as late as 2008, European mushrooms may have included toxicity from the 1986 Chernobyl disaster and continued to be studied.]]

Psilocybin, a naturally occurring chemical in certain psychedelic mushrooms such as Psilocybe cubensis, is being studied for its ability to help people suffering from psychological disorders, such as obsessive–compulsive disorder. Minute amounts have been reported to stop cluster and migraine headaches.

A few species in the genus Amanita, most recognizably A. muscaria, but also A. pantherina, among others, contain the psychoactive compound muscimol. The muscimol-containing chemotaxonomic group of Amanitas contains no amatoxins or phallotoxins, and as such are not hepatoxic, though if not properly cured will be non-lethally neurotoxic due to the presence of ibotenic acid. The Amanita intoxication is similar to Z-drugs in that it includes CNS depressant and sedative-hypnotic effects, but also dissociation and delirium in high doses.

A third type of hallucinogenic mushroom is hallucinogenic bolete mushrooms such as Lanmaoa asiatica, which are said to cause people to experience Lilliputian hallucinations. These mushrooms have unknown constituents and an unknown mechanism of action, but appear to be distinct in their properties and effects from other hallucinogenic mushrooms and are currently being researched.

Folk medicine

thumb|[[Ganoderma lingzhi]]

Some mushrooms are used in folk medicine. In a few countries, extracts, such as polysaccharide-K, schizophyllan, polysaccharide peptide, or lentinan, are government-registered adjuvant cancer therapies, Although some mushroom species or their extracts may be consumed for therapeutic effects, some regulatory agencies, such as the US Food and Drug Administration, regard such use as a dietary supplement, which does not have government approval or common clinical use as a prescription drug.

Some fungi, types of polypores loosely called mushrooms, have been used as fire starters (known as tinder fungi).

Mushrooms and other fungi play a role in the development of new biological remediation techniques (e.g., using mycorrhizae to spur plant growth) and filtration technologies (e.g. using fungi to lower bacterial levels in contaminated water).

<gallery>

File:Amanita muscaria (fly agaric).JPG|Amanita muscaria, a psychotropic mushroom commonly known as "fly agaric"

File:2016-01_Agaricus_bisporus_01.jpg|Agaricus bisporus, a cultivated edible mushroom with various names including "button mushroom", "portobello" and "champignon"

File:Boletus_edulis_02a(js)_Lodz_(Poland).jpg|Boletus edulis, also known as "cep", an edible wild bolete found in Europe

File:Maitake mushroom.jpg|Maitake, a polypore mushroom

File:Zwam, inktzwam.jpg|Coprinopsis atramentaria, commonly known as the "ink cap"

File:Yellowmushrooms.jpg|Leucocoprinus birnbaumii, commonly known as the "flowerpot parasol", at various stages of development

File:Close-up cross section of mushroom.jpg|Close-up cross section of mushroom gills

File:Pop-up mushroom.jpg|Agaricus bitorquis (the "pavement mushroom") emerging through asphalt concrete in summer

</gallery>

See also

  • List of Chinese mushrooms and fungi
  • List of mushroom dishes
  • List of psilocybin mushroom species
  • Largest fungal fruit bodies
  • Lists of fungal species
  • Mushrooms in art

References

Citations

Sources

Identification

  • Mushroom Observer, a collaborative mushroom recording and identification project
  • An Aid to Mushroom Identification , Simon's Rock College
  • Online Edible Wild Mushroom Field Guide