Amushroom ortoadstool is the fleshy,spore-bearingfruiting body of afungus, typically produced above ground, on soil, or on itsfood source.Toadstool generally denotes one poisonous to humans.[1]
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 someAscomycota. The gills produce microscopicspores which help the fungus spread across the ground or its occupant surface.
Forms deviating from the standardmorphology 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 toAgaricus or their orderAgaricales. By extension, the term "mushroom" can also refer to either the entire fungus when in culture, thethallus (calledmycelium) of species forming the fruiting bodies called mushrooms, or the species itself.
Etymology
Amanita muscaria, the most easily recognised "toadstool", is frequently depicted in fairy stories and on greeting cards. It is often associated withgnomes.[2]
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 termsmushrom, mushrum, muscheron, mousheroms, mussheron, or musserouns were used.[3]
The term "mushroom" and its variations may have been derived from the French wordmousseron in reference tomoss (mousse). Delineation between edible and poisonous fungi is not clear-cut, so a "mushroom" may be edible, poisonous, or unpalatable.[4][5] The wordtoadstool appeared first in14th century England as a reference for a "stool" fortoads, possibly implying an inedible poisonous fungus.[6]
Identification
Identifying what is and is not a mushroom requires a basic understanding of theirmacroscopic structure. Most arebasidiomycetes and gilled. Their spores, calledbasidiospores, 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 offbasidia 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 aspore 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.[7]
Morphological characteristics of the caps of mushrooms
While modern identification of mushrooms is quickly becoming molecular, the standard methods for identification are still used by most and have developed into a fine art harking back tomedieval times and theVictorian era, combined with microscopic examination. The presence of juices upon breaking, bruising-reactions, odors, tastes, shades of color, habitat, habit, and season are all considered by both amateur and professional mycologists. Tasting and smelling mushrooms carries its own hazards because of poisons andallergens.Chemical tests are also used for some genera.[8]
In general, identification togenus can often be accomplished in the field using a localfield guide. Identification tospecies, 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 onlamella edges for colored spored prints.
An atypical mushroom is thelobster mushroom, which is a fruitbody of aRussula orLactarius mushroom that has been deformed by theparasitic fungusHypomyces lactifluorum. This gives the affected mushroom an unusual shape and red color that resembles that of a boiledlobster.[9]
Other mushrooms are not gilled, so the term "mushroom" is loosely used, and giving a full account of their classifications is difficult. Some have pores underneath (and are usually calledboletes), others have spines, such as thehedgehog mushroom and othertooth fungi, and so on. "Mushroom" has been used forpolypores,puffballs,jelly fungi,coral fungi,bracket fungi,stinkhorns, andcup fungi. Thus, the term is more one of common application tomacroscopic fungal fruiting bodies than one having precisetaxonomic meaning. Approximately 14,000 species of mushrooms are described.[10]
A mushroom develops from a nodule, or pinhead, less than two millimeters in diameter, called aprimordium, which is typically found on or near the surface of thesubstrate. It is formed within themycelium, the mass of threadlikehyphae that make up the fungus. The primordium enlarges into a roundish structure of interwoven hyphae roughly resembling an egg, called a "button". The button has a cottony roll of mycelium, theuniversal veil, that surrounds the developing fruit body. As the egg expands, the universal veil ruptures and may remain as a cup, orvolva, at the base of thestalk, or as warts or volval patches on the cap. Many mushrooms lack a universal veil, therefore they do not have either a volva or volval patches. Often, a second layer of tissue, thepartial veil, covers the bladelikegills that bearspores. As the cap expands the veil breaks, and remnants of the partial veil may remain as a ring, orannulus, around the middle of the stalk or as fragments hanging from the margin of the cap. The ring may be skirt-like as in some species ofAmanita, collar-like as in many species ofLepiota, or merely the faint remnants of a cortina (a partial veil composed of filaments resembling a spiderweb), which is typical of the genusCortinarius. Mushrooms lacking partial veils do not form an annulus.[11]
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 ofPleurotus andPanus. 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 includingtruffles,jellies,earthstars, andbird'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 generaAgaricus,Amanita,Lepiota andPluteus, among others, have free gills that do not extend to the top of the stalk. Others havedecurrent gills that extend down the stalk, as in the generaOmphalotus andPleurotus. 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.[12]
Microscopic features
Ahymenium 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 ofboletes 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 calledasci, which typically contain eight spores in each ascus. TheDiscomycetes, which contain the cup, sponge, brain, and some club-like fungi, develop an exposed layer of asci, as on the inner surfaces ofcup fungi or within the pits ofmorels. ThePyrenomycetes, 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 calledperithecia, within which the asci develop.[13]
In the basidiomycetes, usually four spores develop on the tips of thin projections calledsterigmata, which extend from club-shaped cells called abasidia. The fertile portion of theGasteromycetes, called agleba, may become powdery as in the puffballs or slimy as in thestinkhorns. Interspersed among the asci are threadlike sterile cells calledparaphyses. Similar structures calledcystidia 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.[13]
The most important microscopic feature for identification of mushrooms is the spores. Their color, shape, size, attachment, ornamentation, and reaction tochemical tests often can be the crux of an identification. A spore often has a protrusion at one end, called an apiculus, which is the point of attachment to the basidium, termed the apicalgerm pore, from which the hypha emerges when the spore germinates.[13]
Many species of mushrooms seemingly appear overnight, growing or expanding rapidly. This phenomenon is the source of several common expressions in theEnglish language including "to mushroom" or "mushrooming" (expanding rapidly in size or scope) and "to pop up like a mushroom" (to appear unexpectedly and quickly). In reality, all species of mushrooms take several days to form primordial mushroom fruit bodies, though they do expand rapidly by the absorption of fluids.[14][15][16][17]
Thecultivated mushroom, as well as the commonfield mushroom, initially form a minutefruiting 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 itsmycelium and expand, mainly by inflating preformedcells that took several days to form in theprimordia.[18]
Similarly, there are other mushrooms, likeParasola plicatilis (formerlyCoprinus plicatlis), that grow rapidly overnight and may disappear by late afternoon on a hot day after rainfall.[19] The primordia form at ground level in lawns in humid spaces under thethatch and after heavy rainfall or indewy conditions balloon to full size in a few hours, release spores, and then collapse.[20][21]
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 insertinghyphae. For example,Pleurotus nebrodensis grows slowly, and because of this combined with human collection, it is nowcritically endangered.[22]
Though mushroom fruiting bodies are short-lived, the underlying mycelium can itself be long-lived and massive. A colony ofArmillaria solidipes (formerly known asArmillaria ostoyae) inMalheur National Forest in theUnited States is estimated to be 2,400 years old, possibly older, and spans an estimated 2,200 acres (8.9 km2).[23] Most of the fungus is underground and in decaying wood or dying tree roots in the form of white mycelia combined with black shoelace-likerhizomorphs that bridge colonized separated woody substrates.[24]
Thevitamin D content of a mushroom depends onpostharvest handling, in particular the unintended exposure to sunlight. TheUS Department of Agriculture provided evidence that UV-exposed mushrooms contain substantial amounts of vitamin D.[27] When exposed toultraviolet (UV) light, even after harvesting,[28]ergosterol in mushrooms is converted tovitamin D2,[29] a process now used intentionally to supply fresh vitamin D mushrooms for thefunctional foodgrocery market.[30][31] 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 naturalsunlight, and that UV light has a long record of safe use for production of vitamin D in food.[30]
Most mushrooms sold insupermarkets have been commercially grown onmushroom 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 ofA. bisporus are grown commercially, including whites, crimini, and portobello. Other cultivated species available at many grocers includeHericium erinaceus,shiitake,maitake (hen-of-the-woods),Pleurotus, andenoki. 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.[33]
China is a major edible mushroom producer.[34] The country produces about half of all cultivated mushrooms, and around 2.7 kilograms (6.0 lb) of mushrooms are consumed per person per year by 1.4 billion people.[35] In 2014,Poland was the world's largest mushroom exporter, reporting an estimated 194,000 tonnes (191,000 long tons; 214,000 short tons) annually.[36]
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,[37] and the act of collecting them for such is known asmushroom hunting, or simply "mushrooming". Even edible mushrooms may produceallergic reactions in susceptible individuals, from a mildasthmatic response to severeanaphylactic shock.[38][39] Even the cultivatedA. bisporus contains small amounts ofhydrazines, the most abundant of which isagaritine (amycotoxin andcarcinogen).[40] However, the hydrazines are destroyed by moderate heat when cooking.[41]
A number of species of mushrooms arepoisonous; although some resemble certain edible species, consuming them could be fatal. Eating mushrooms gathered in the wild is risky and should only be undertaken by individuals knowledgeable in mushroom identification. Common best practice is for wild mushroom pickers to focus on collecting a small number of visually distinctive, edible mushroom species that cannot be easily confused with poisonous varieties. Common mushroom hunting advice is that if a mushroom cannot be positively identified, it should be considered poisonous and not eaten.[42]
YoungAmanita phalloides "death cap" mushrooms, with a matchbox for size comparison
Many mushroom species producesecondary metabolites that can be toxic, mind-altering, antibiotic, antiviral, orbioluminescent. Although there are only a small number ofdeadly 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 (seeemetics), or to learn to avoid consumption altogether. In addition, due to the propensity of mushrooms to absorbheavy metals, including those that are radioactive, as late as 2008, European mushrooms may have included toxicity from the 1986Chernobyl disaster and continued to be studied.[43][44]
Mushrooms with psychoactive properties have long played a role in various native medicine traditions in cultures all around the world. They have been used as sacrament in rituals aimed at mental and physical healing, and to facilitate visionary states. One such ritual is thevelada ceremony. A practitioner of traditional mushroom use is theshaman orcurandera (priest-healer).[45]
Psilocybin mushrooms, also referred to as psychedelic mushrooms, possesspsychedelic properties. Commonly known as "magic mushrooms" or"'shrooms", they are openly available insmart shops in many parts of the world, or on theblack market in those countries which have outlawed their sale. Psilocybin mushrooms have been reported to facilitate profound and life-changing insights often described asmystical experiences. Recent scientific work has supported these claims, as well as the long-lasting effects of such induced spiritual experiences.[46]
There are over 100 psychoactive mushroom species of genusPsilocybe native to regions all around the world.[47]
Psilocybin, a naturally occurring chemical in certainpsychedelic mushrooms such asPsilocybe cubensis, is being studied for its ability to help people suffering from psychological disorders, such asobsessive–compulsive disorder. Minute amounts have been reported to stopcluster andmigraine headaches.[48] A double-blind study, done byJohns Hopkins Hospital, showed psychedelic mushrooms could provide people an experience with substantial personal meaning and spiritual significance. In the study, one third of the subjects reported ingestion of psychedelic mushrooms was the single most spiritually significant event of their lives. Over two-thirds reported it among their five most meaningful and spiritually significant events. On the other hand, one-third of the subjects reported extremeanxiety. However the anxiety went away after a short period of time.[49] Psilocybin mushrooms have also shown to be successful in treating addiction, specifically with alcohol and cigarettes.[50]
Mushrooms can be used fordyeingwool and other natural fibers. Thechromophores ofmushroom dyes are organic compounds and produce strong and vivid colors, and all colors of the spectrum can be achieved with mushroom dyes. Before the invention of syntheticdyes, mushrooms were the source of many textile dyes.[56]
Some fungi, types ofpolypores loosely called mushrooms, have been used as fire starters (known astinder fungi).
Mushrooms and other fungi play a role in the development of new biological remediation techniques (e.g., usingmycorrhizae to spur plant growth) and filtration technologies (e.g. using fungi to lower bacterial levels in contaminated water).[57]
There is an ongoing research in the field of genetic engineering aimed towards creation of the enhanced qualities of mushrooms for such domains as nutritional value enhancement, as well as medical use.[58]
Gallery
Amanita muscaria, a psychotropic mushroom commonly known as "fly agaric"
Agaricus bisporus, a cultivated edible mushroom with various names including "button mushroom", "portobello" and "champignon"
^Falconer, William (2009).Mushrooms: How to Grow Them - A Practical Treatise on Mushroom Culture for Profit and Pleasure (2nd ed.). Read Books.ISBN9781444678925.
^Gordon, Tom (2021).A Complete Mushroom Cultivation Guide on How to Grow Gourmet Mushrooms and Identify Wild Common Mushrooms and Other Fungi for Beginners. Independently Published.ISBN9798702942391.
^Haytowitz DB (2009)."Vitamin D in mushrooms"(PDF). Nutrient Data Laboratory, US Department of Agriculture.Archived(PDF) from the original on 1 February 2021. Retrieved16 April 2018.
^Kalaras, M. D.; Beelman, R. B.; Elias, R. J. (2012). "Effects of postharvest pulsed UV light treatment of white button mushrooms (Agaricus bisporus) on vitamin D2 content and quality attributes".Journal of Agricultural and Food Chemistry.60 (1):220–5.Bibcode:2012JAFC...60..220K.doi:10.1021/jf203825e.PMID22132934.
^abSimon, R. R.; Borzelleca, J. F.; Deluca, H. F.; Weaver, C. M. (2013). "Safety assessment of the post-harvest treatment of button mushrooms (Agaricus bisporus) using ultraviolet light".Food and Chemical Toxicology.56:278–89.doi:10.1016/j.fct.2013.02.009.PMID23485617.
^Siegered, Agnes A., ed. (January 1998)."Spore Prints #338".Bulletin of the Puget Sound Mycological Society.Archived from the original on 17 July 2010. Retrieved4 July 2010.
^Griffiths RR, Richards WA, McCann U, Jesse R (2006). "Psilocybin can occasion mystical-type experiences having substantial and sustained personal meaning and spiritual significance".Psychopharmacology.187 (3):268–283.doi:10.1007/s00213-006-0457-5.PMID16826400.S2CID7845214.
Ammirati JF, Traquair JA, Horgen PA (1985).Poisonous Mushrooms of Canada: Including other Inedible Fungi. Markham, Ontario: Fitzhenry & Whiteside in cooperation with Agriculture Canada and the Canadian Government Publishing Centre, Supply and Services Canada.ISBN978-0-88902-977-4.
Hall IR, Stephenson SL, Buchanan PK, Yun W, Cole AL (2003).Edible and Poisonous Mushrooms of the World. Portland, Oregon: Timber Press.ISBN978-0-88192-586-9.
Stuntz DE, Largent DL, Thiers HD, Johnson DJ, Watling R (1978).How to Identify Mushrooms to Genus I. Eureka, California: Mad River Press.ISBN978-0-916422-00-4.
External links
Look upmushroom in Wiktionary, the free dictionary.
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