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''Morchella esculenta'' (commonly known as common morel, morel, yellow morel, true morel, morel mushroom, and sponge morel) is a species of
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the family Morchellaceae of the
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
. It is one of the most readily recognized of all the
edible mushroom Edible mushrooms are the fleshy fruit bodies of numerous species of macrofungi (fungi that bear fruiting structures large enough to be seen with the naked eye). Edibility may be defined by criteria including the absence of poisonous effect ...
s and highly sought after. Each fruit body begins as a tightly compressed, grayish sponge with lighter ridges, and expands to form a large yellowish sponge with large pits and ridges raised on a large white stem. The pitted yellow-brown caps measure broad by tall, and are fused to the stem at its lower margin, forming a continuous hollow. The pits are rounded and irregularly arranged. The hollow stem is typically long by thick, and white to yellow. The fungus fruits under hardwoods and conifers during a short period in the spring, depending on the weather, and is also associated with old orchards, woods and disturbed grounds.


Description

The
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. The origin of the word "cap" comes from the Old French word "chapeau" which means "head co ...
is pale brownish cream, yellow to tan or pale brown to grayish brown. The edges of the ridges are usually lighter than the pits, and somewhat oval in outline, sometimes bluntly cone-shaped with a rounded top or more elongate. Caps are hollow, attached to the stem at the lower edge, and typically about broad by tall. The flesh is brittle. The stem is white to pallid or pale yellow, hollow, and straight or with a club-shaped or bulbous base. It is finely granular overall, somewhat ridged, generally about long by thick. In age it may have brownish stains near the base. It has a passing resemblance to the common stinkhorn ('' Phallus impudicus''), for which it is sometimes mistaken. Yellow morels are often found near wooded areas. Centipedes sometimes make their home inside these morels; infested morels usually have a hole in the top.


Microscopic characteristics

The
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s range from white to cream to slightly yellow in deposit, although a spore print may be difficult to obtain given the shape of the fruit body. The spores are formed in asci lining the pits—the ridges are sterile. They are ellipsoidal, smooth, thin-walled, translucent (
hyaline A hyaline substance is one with a glassy appearance. The word is derived from , and . Histopathology Hyaline cartilage is named after its glassy appearance on fresh gross pathology. On light microscopy of H&E stained slides, the extracellula ...
), and measure 17.5–21.9 by 8.8–11.0  μm. The asci are eight-spored, 223–300 by 19–20 μm, cylindrical, and hyaline. The paraphyses are filamentous, cylindrical, 5.8–8.8 μm wide, and hyaline. The
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e of the stem are interwoven, hyaline, and measure 5.8–9.4 μm wide. The surface hyphae are inflated, spherical to pear-shaped, 22–44 μm wide, covered by a network of interwoven hyphae 11–16.8 μm wide with recurved cylindrical hyphal ends.


Development

Fruit bodies have successfully been grown in the laboratory. R. Ower was the first to describe the developmental stages of ascomata grown in a controlled chamber. This was followed by in-depth cytological studies by Thomas Volk and Leonard (1989, 1990). To study the morel life cycle they followed the development of ascoma fruiting in association with tuberous begonias ('' Begonia tuberhybrida''), from very small primordia to fully developed fruit bodies. Young fruit bodies begin development in the form of a dense knot of hyphae, when suitable conditions of moisture and nutrient availability conditions have been reached. Hyphal knots are underground and cup-shaped for some time, but later emerge from the soil and develop into a stalked fruiting body. Further growth makes the
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
convex with the asci facing towards the outer side. Because of the unequal growth of the surface of the hymenium, it becomes folded to form many ridges and depressions, resulting in the sponge or honeycomb appearance.


Similar species

''Morchella esculenta'' is probably the most familiar of the morels. In contrast to '' M. angusticeps'' and its relatives, the caps are light-colored throughout development, especially the ridges, which remain paler than the pits. '' M. crassipes'' is sometimes confused with ''M. esculenta''. According to Smith (1975), the two are distinct, but young forms of ''M. crassipes'' are difficult to separate from ''M. esculenta''. The two are similar in color, but ''M. crassipes'' is larger, often has thin ridges, and sometimes has a stem base that is enlarged and longitudinally grooved. Stinkhorns (esp. ''Phallus impudicus'') have also been confused with morels, but specimens of the former have a volva at the base of the stem, and are covered with gleba—a slimy, foul-smelling spore mass. See also ''
Gyromitra esculenta ''Gyromitra esculenta'' is an ascomycete fungus from the genus ''Gyromitra'', widely distributed across Europe and North America. It normally fruits in sandy soils under coniferous trees in spring and early summer. The Ascocarp, fruiting bod ...
'', which is similar in appearance but poisonous.


Taxonomy

The fungus was originally named ''Phallus esculentus'' by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in his ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genus, genera. It is the first work to consistently apply binomial nomenclature ...
'' (1753), and given its current name by Swedish mycologist
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. He is sometimes called the Mycology, "Linnaeus of Mycology". In his works he described and assigned botanical names to hundreds of fungus and li ...
in 1801. ''Morchella esculenta'' is commonly known by various names: morel, common morel, true morel, morel mushroom, yellow morel, sponge morel, Molly Moocher, haystack, and dryland fish. In
Nepal Nepal, officially the Federal Democratic Republic of Nepal, is a landlocked country in South Asia. It is mainly situated in the Himalayas, but also includes parts of the Indo-Gangetic Plain. It borders the Tibet Autonomous Region of China Ch ...
it is known as ''Guchi chyau''. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
is derived from the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''esculenta'', meaning "edible". The scientific name ''Morchella esculenta'' has been applied to many similar yellow morels throughout the world. In 2014 Richard et al. used DNA analysis to restrict the ''M. esculenta'' name to a single species of yellow morel commonly found in Europe and also reported from China. Other species of yellow morel, including those in North America, have received new scientific names.


Distribution and habitat

Fruit bodies are sometimes solitary, but more often in groups, on the ground in a variety of habitats. A preference for soil with a
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
base (alkaline) has been noted, but they have also been found in acid soils. The mushroom is usually found in early spring, in forests, orchards, yards, gardens and sometimes in recently burned areas. In North America, it is sometimes referred to as the "May mushroom" due to its consistent fruiting in that month, but the time of fruiting varies locally, from February to July. It is typically the last morel species to fruit in locales where more than one species are found. For example, in northern Canada and in cooler mountainous regions, morels typically do not appear until June. It has been suggested that the springtime fruiting may be due to their ability to grow at low temperatures to the exclusion of competition, a conclusion later corroborated by experiments correlating spore
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ...
to soil temperatures. One author suggests the acronym PETSBASH may be used to remember the trees associated with morels:
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
, elm,
tulip Tulips are spring-blooming perennial herbaceous bulbiferous geophytes in the ''Tulipa'' genus. Their flowers are usually large, showy, and brightly coloured, generally red, orange, pink, yellow, or white. They often have a different colour ...
, sassafras,
beech Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
, ash, sycamore, and
hickory Hickory is a common name for trees composing the genus ''Carya'', which includes 19 species accepted by ''Plants of the World Online''. Seven species are native to southeast Asia in China, Indochina, and northeastern India (Assam), and twelve ...
. In North America, it is widely distributed, but especially common in eastern North America and the Midwest. David Arora notes that "large crops can also be found around the bases of dying (but not quite dead) elms attacked by Dutch elm disease." The species has been named state mushroom of
Minnesota Minnesota ( ) is a U.S. state, state in the Upper Midwestern region of the United States. It is bordered by the Canadian provinces of Manitoba and Ontario to the north and east and by the U.S. states of Wisconsin to the east, Iowa to the so ...
, and was the first state mushroom of any state. It can also be found in Brazil and Bulgaria. In Jammu and Kashmir wild mushrooms, locally known as Himalayan wild mushroom, Gucchi, '' Morchella conica'' and ''Morchella esculenta'', are gathered and supplied as medical remedy.


Cultivation

Due to the mushroom's prized fruit bodies, several attempts have been made to grow the fungus in culture. In 1901, Repin reported successfully obtaining fruit bodies in a cave in which cultures had been established in flower pots nine years previously in 1892. Mycologist Taylor Piercefield developed a method using beds of hardwood tree saplings which would be inoculated with mycelium, concentrating on th
symbiotic relationship of Morchella esculenta
Later, once the mycelium network had been fully developed, the beds would be treated with potassium hydroxide to replicate the pH conditions found in soil after a forest fire. This method resulted in large, mature fruits, but was not commercially viable on a large scale. More recently, small scale commercial growers have had success growing morels by using partially shaded rows of mulched wood. The rows of mulch piles are inoculated with morel mushroom spores in a solution of water and molasses which are poured over the piles of mulch and then they are allowed to grow undisturbed for several weeks. A solution of wood ashes mixed in water and diluted is subsequently poured over the rows of wood mulch which triggers fruiting of the morels. Morels are known to appear after fires and the alkaline conditions produced by wood ash mixed with water initiate fruit body formation for most species of morels.


Uses


Edibility

''Morchella esculenta'', like all morels, are among the most highly prized of all
edible mushroom Edible mushrooms are the fleshy fruit bodies of numerous species of macrofungi (fungi that bear fruiting structures large enough to be seen with the naked eye). Edibility may be defined by criteria including the absence of poisonous effect ...
s. Raw morels have a gastrointestinal irritant,
hydrazine Hydrazine is an inorganic compound with the chemical formula . It is a simple pnictogen hydride, and is a colourless flammable liquid with an ammonia-like odour. Hydrazine is highly hazardous unless handled in solution as, for example, hydraz ...
(this has never been found in morel samples however it is assumed), but parboiling or blanching before consumption will remove it. Old fruit bodies that show signs of decay may be poisonous. The mushrooms may be fried in butter or baked after being stuffed with meats and vegetables. The mushrooms may also be dried by threading the caps onto string and hanging them in the sun; this process is said to concentrate the flavour. One study determined the main nutritional components to be as follows (on a dry weight basis):
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
32.7%,
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers specif ...
2.0%,
fiber Fiber (spelled fibre in British English; from ) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often inco ...
17.6%, ash 9.7%, and
carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...
s 38.0%.


Bioactive compounds

Both the fruit bodies and the mycelia of ''M. esculenta'' contain an uncommon
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...
, ''cis''-3-amino-L-proline; this amino acid does not appear to be protein bound. In addition to ''M. esculenta'', the amino acid is known to exist only in '' M. conica'' and '' M. crassipes.''


Industrial applications; solid-state fermentation

Solid-state fermentation is an industrial process to produce
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s and to upgrade the values of existing foods, especially foods from East Asia. Solid-state fermentation is a process whereby an insoluble substrate is fermented with sufficient moisture but without free water. Solid-state fermentation, unlike that of slurry state, requires no complex fermentation controls and has many advantages over submerged liquid fermentation. ''M. esculenta'' has shown promise in degrading
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diet ...
and upgrading the nutritional value of cornmeal during solid-state fermentation. ''M. esculenta'' mycelia is able to bind to and inhibit the effects of furanocoumarins, chemicals found in
grapefruit The grapefruit (''Citrus'' × ''paradisi'') is a subtropical citrus tree known for its relatively large, sour to semi-sweet, somewhat bitter fruit. The flesh of the fruit is segmented and varies in color from pale yellow to dark red. Grapefru ...
that inhibit human
cytochrome p450 Cytochromes P450 (P450s or CYPs) are a Protein superfamily, superfamily of enzymes containing heme as a cofactor (biochemistry), cofactor that mostly, but not exclusively, function as monooxygenases. However, they are not omnipresent; for examp ...
enzymes and are responsible for the "grapefruit/drug" interaction phenomenon.


References


External links

* {{Taxonbar, from=Q1129282 esculenta Fungi of Europe Fungi of North America Edible fungi Taxa named by Carl Linnaeus Fungi described in 1753 Fungus species