Morchella esculenta

Morchella esculenta
Scientific classification
Kingdom:	Fungi
Division:	Ascomycota
Subdivision:	Pezizomycotina
Class:	Pezizomycetes
Order:	Pezizales
Family:	Morchellaceae
Genus:	Morchella
Species:	M. esculenta
Binomial name
Morchella esculenta Fr.
Synonyms
Helvella esculenta (L.) Sowerby Phallus esculentus L.

Morchella esculenta
Mycological characteristics
	smooth hymenium
	cap is conical or ovate
	hymenium attachment is irregular or not applicable
	stipe is bare
	spore print is cream to yellow
	ecology is mycorrhizal or saprotrophic
	edibility: choice

Morchella esculenta, (commonly known as common morel, morel, yellow morel, true morel, morel mushroom, and sponge morel) is a species of fungus in the Morchellaceae family of the Ascomycota. It is one of the most readily recognized of all the edible mushrooms 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 2–7 cm (0.8–2.8 in) broad by 2–10 cm (0.8–3.9 in) 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 2–9 cm (0.8–3.5 in) long by 2–5 cm (0.8–2.0 in) thick, and white to yellow. The fungus fruits under hardwoods during a short period in the spring, depending on the weather, but it is also associated with old orchards, woods, disturbed grounds and burnt areas. Although a process was reported in 1982 to grow the fruit bodies under controlled conditions, attempts to cultivate the mushroom commercially have only been partially successful.

Taxonomy and naming

The fungus was originally named Phallus esculentus by Carl Linnaeus in his Species Plantarum (1753),^[1] and given its current name by Swedish mycologist Elias Magnus Fries in 1801.^[2][3]

Morchella esculenta is commonly known by various names: morel, common morel, true morel, morel mushroom, yellow morel, sponge morel,^[4] Molly Moocher, haystack, and dryland fish.^[5] In Nepal it is known as Guchi chyau.^[6] The specific epithet is derived from the Latin esculenta, meaning "edible".

Description

The cap is pale brownish cream, yellow to tan or pale brown to grayish brown. The edges of the ridges are usually not darker than the pits, and somewhat oval in outline, sometimes bluntly cone-shaped with a rounded top or more elongate. Caps are hollow, and attached to the stem at the lower edge, and typically about 2–7 cm (0.8–2.8 in) broad by 2–10 cm (0.8–3.9 in) 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 2–9 cm (0.8–3.5 in) long by 2–5 cm (0.8–2.0 in) thick.^[7] In age it may have brownish stains near the base.^[5]

Microscopic characteristics

The spores 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.^[8] The spores are formed in asci lining the pits—the ridges are sterile.^[9] They are ellipsoidal, smooth, thin-walled, translucent (hyaline), 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.^[10]

The hyphae 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 described the developmental stages of ascomata grown in a controlled chamber.^[11] 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.^[12][13]

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 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.^[14]

Similar species

Morchella esculenta is probably the most popular 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.^[10]

Morels have also been confused with stinkhorns,^[15] but specimens of the latter have a volva at the base of the stem, are covered with gleba—a slimy, foul-smelling spore mass.

Habitat and distribution

Drawing by von Albin Schmalfuß, 1897

Fruit bodies are sometimes found solitary, but more often in groups, on the ground in a variety of habitats. A preference for soil with a limestone base (alkaline) has been noted,^[5] but they have also been found in acid soils.^[16] The mushroom is usually found in early spring, in forests, orchards, yards, gardens and sometimes in recently burned areas.^[7] 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.^[17] For example, in Northern Canada and in cooler mountainous regions, morels typically do not appear until June.^[18] It has been suggested that the springtime fruiting may be due to their ability to grow at low temperatures to the exclusion of competition,^[19] a conclusion later corroborated by experiments correlating spore germination to soil temperatures.^[20]

One author suggests the acronym PETSBASH may be used to remember the trees associated with morels: pine, elm, tulip, sassafras, beech, ash, sycamore, and hickory.^[21]

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."^[15] The species has been named state mushroom of Minnesota, and was the first state mushroom of any state.^[22][23]

Brazil^[24]

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.^[25]

Uses

Edibility

Harvested morels

Morchella esculenta, like all morels, are among the most highly-prized of all edible mushrooms. Raw mushrooms have a gastrointestinal irritant, but parboiling or blanching before consumption will remove it. Old fruit bodies that show signs of decay may be poisonous.^[8] The mushrooms may be fried in butter or baked after being stuffed with meats and vegetables.^[26] 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 flavor.^[8] One study determined the main nutritional components to be as follows (on a dry weight basis): protein 32.7%, fat 2.0%, fiber 17.6%, ash 9.7%, and carbohydrates 38.0%.^[27]

In one isolated case in Germany, six people were reported to have developed neurologic effects between 6–12 hours after consumption. The effects included ataxia and visual disturbances, and lasted up to a day before disappearing without enduring effects.^[28]

Bioactive compounds

Both the fruit bodies and the mycelia of M. esculenta contain an uncommon amino acid, cis-3-amino-L-proline; this amino acid does not appear to be protein bound.^[29] In addition to M. esculento, the amino acid is known to exist only in M. conica and M. crassipes.^[30]

Medicinal properties

Laboratory experiments using rodent models suggest that the polysaccharides from M. esculenta fruit bodies have several medicinal properties, including anti-tumor effects, immunoregulatory properties,^[31] fatigue resistance, and antiviral effects.^[32][33][34] Extracts from the fruit bodies have antioxidant properties.^[35] It also has been shown that the polysaccharides from M. esculenta mycelia have antioxidant activity.^[36][37] The fungus is listed in the IUCN National Register of medicinal plants in Nepal.^[6]

Morchella esculenta is also used in traditional Chinese medicine to treat indigestion, excessive phlegm, and shortness of breath.^[38]

Industrial applications; solid state-fermentation

Solid-state fermentation (SSF) is an industrial process to produce enzymes and to upgrade the values of existing foods, especially oriental foods. SSF is a process whereby an insoluble substrate is fermented with sufficient moisture but without free water. SSF, 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 and upgrading the nutritional value of cornmeal during SSF.^[39]

M. esculenta mycelia is able to bind to and inhibit the effects of Furanocoumarins, chemicals found in grapefruit that inhibit human cytochrome p450 enzymes and are responsible for the "grapefruit/drug" interaction phenomenon.^[40]

References

1. ^ Fries EM. (1753) (in Latin). Species Plantarum. pp. 1178–79. http://gallica.bnf.fr/ark:/12148/bpt6k96633w.image.f620.pagination. Retrieved 2010-03-21. 
2. ^ Persoon CH. (1801). Synopsis Methodica Fungorum. 2. p. 618. http://194.203.77.76/LibriFungorum/Image.asp?ItemID=82&ImageFileName=0618b.jpg. Retrieved 2010-03-21. 
3. ^ Kuo M. "Morchella esculenta". MushroomExpert.Com. http://www.mushroomexpert.com/morchella_esculenta.html. Retrieved 2010-03-26. 
4. ^ Institut Fur Pflanzengenetik Und Kulturpflanzenforschung Gatersleben, Hanelt P, Buttner R, Rudolf Vorlaufiges Verzeichnis Landwirtschaftlich Oder gart Mansfeld (2001). Mansfeld's Encyclopedia of Agricultural and Horticultural Crops: (Except Ornamentals). Santa Clara, California: Springer-Verlag TELOS. p. 17. ISBN 3-540-41017-1. http://books.google.com/?id=10IMFSavIMsC&lpg=PA17&dq=Morchella%20esculenta&pg=PA17#v=onepage&q=Morchella%20esculenta&f=false. Retrieved 2010-03-21. 
5. ^ ^{a b c} Roody WC. (2003). Mushrooms of West Virginia and the Central Appalachians. Lexington, Kentucky: University Press of Kentucky. p. 485. ISBN 0-8131-9039-8. http://books.google.com/?id=5HGMPEiy4ykC&lpg=PA485&dq=morchella%20esculenta&pg=PA485#v=onepage&q=morchella%20esculenta&f=false. Retrieved 2010-03-22. 
6. ^ ^{a b} National register of medicinal plants. IUCN-the World Conservation Union: His Majesty's Government, Ministry of Forest and Soil Conservation. 2000. p. 61. ISBN 978-9291440481. http://books.google.com/?id=CgHsWXkgbX0C&lpg=PA61&dq=Morchella%20esculenta&pg=PA61#v=onepage&q=Morchella%20esculenta&f=false. Retrieved 2010-03-21. 
7. ^ ^{a b} Ammirati JF, McKenny M, Stuntz DE. (1987). The New Savory Wild Mushroom. Seattle: University of Washington Press. pp. 209–10. ISBN 0-295-96480-4. 
8. ^ ^{a b c} Hall IR. (2003). Edible and Poisonous Mushrooms of the World. Portland, Or: Timber Press. pp. 239–42. ISBN 0-88192-586-1. 
9. ^ Schalkwijk-Barendsen HME. (1991). Mushrooms of Western Canada. Edmonton: Lone Pine Publishing. pp. 381–82. ISBN 0-919433-47-2. 
10. ^ ^{a b} Ammirati J, Traquair JA, Horgen PA. (1985). Poisonous Mushrooms of Canada. Fitzhenry & Whiteside in cooperation with Agriculture Canada. pp. 287–88. ISBN 978-0889029774. 
11. ^ Ower R. (1982). "Notes on the development of the morel ascocarp: Morchella esculenta". Mycologia 74 (1): 142–44. doi:10.2307/3792639. JSTOR 3792639. 
12. ^ Volk TJ, Leonard TJ. (1989). "Experimental studies on the morel. I. Hetrokaryon formation between mono ascosporous strains of Morchella". Mycologia 81 (4): 523–31. doi:10.2307/3760127. JSTOR 3760127. 
13. ^ Volk T, Leonard T. (1990). "Cytology of the life-cycle of Morchella'". Mycological Research 94: 399–406. doi:10.1016/S0953-7562(09)80365-1. 
14. ^ Sharma OP. (1988). Textbook of Fungi. Boston: McGraw Hill Higher Education. pp. 193–96. ISBN 0-07-460329-9. 
15. ^ ^{a b} Arora D. (1986). Mushrooms Demystified: a Comprehensive Guide to the Fleshy Fungi. Berkeley, California: Ten Speed Press. pp. 787–88. ISBN 0-89815-169-4. http://books.google.com/?id=86tM01VsFG0C&lpg=RA1-PA787&dq=morchella%20esculenta&pg=RA1-PA788#v=onepage&q=morchella%20esculenta&f=false. Retrieved 2010-03-22. 
16. ^ Metzler V, Metzler S. (1992). Texas Mushrooms: a Field Guide. Austin: University of Texas Press. p. 330. ISBN 0-292-75125-7. http://books.google.com/?id=HRtfvVigMmsC&pg=PA330&dq=urnula+craterium. Retrieved 2010-03-26. 
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18. ^ Bessette A, Fischer DH. (1992). Edible Wild Mushrooms of North America: a Field-to-Kitchen Guide. Austin: University of Texas Press. pp. 134–35. ISBN 0-292-72080-7. http://books.google.com/?id=zsJN7NZutTAC&lpg=PA134&dq=morchella%20esculenta&pg=PA135#v=onepage&q=morchella%20esculenta&f=false. Retrieved 2010-03-22. 
19. ^ Baker KF, Cook RJ. (1974). Biological control of plant pathogens. San Francisco: W. H. Freeman. ISBN 0716705893. 
20. ^ Schmidt EL. (1983). "Spore germination of and carbohydrate colonization by Morchella esculenta at different soil temperatures". Mycologia 75 (5): 870–75. doi:10.2307/3792778. JSTOR 3792778. 
21. ^ Rosen S. (1982). A Judge Judges Mushrooms. Highlander Pr. ISBN 0-913617-01-6. 
22. ^ "Minnesota State Symbols: Minnesota State Mushroom". Minnesota Legislature. http://www.leg.state.mn.us/leg/symbols.aspx#mushroom. Retrieved June 13, 2011. 
23. ^ "2010 Minnesota Statutes: 1.149 State Mushroom". Minnesota Office of the Revisor of Statutes. https://www.revisor.mn.gov/statutes/?id=1.149. Retrieved June 13, 2011. 
24. ^ Cortez VG, Coelho G, Guerrero RT. (2004). "Morchella esculenta (Ascomycota): A rare species found in Santa Maria, Rio Grande do Sul, Brazil" (in Portuguese). Biociencias (Porto Alegre) 12 (1): 51–53. 
25. ^ Repin C. (1901). "Sur la culture de le Morille". Revue générale des sciences pures et appliquées 12: 595–96. http://www.biodiversitylibrary.org/item/41549#605. Retrieved 2010-03-21. 
26. ^ Abel D, Horn B, Kay R. (1993). A Guide to Kansas Mushrooms. Lawrence: University Press of Kansas. p. 63. ISBN 0-7006-0571-1. 
27. ^ Wahid M, Sattar A, Khan S. (1988). "Composition of wild and cultivated mushrooms of Pakistan". Mushroom Journal for the Tropics 8 (2): 47–51. 
28. ^ Pfab R, Haberl B, Kleber J, Zilker T. (2008). "Cerebellar effects after consumption of edible morels (Morchella conica, Morchella esculenta)". Clinical Toxicology 46 (3): 259–60. doi:10.1080/15563650701206715. PMID 18344109. 
29. ^ Hatanaka S-I. (1969). "A new amino acid isolated from Morchella esculenta and related species". Phytochemistry 8 (7): 1305–08. doi:10.1016/S0031-9422(00)85571-5. 
30. ^ Moriguchi M, Sada S-I, Hatanaka S-I. (1979). "Isolation of cis-3-amino-L-proline from cultered mycelia or Morchella esculenta". Applied and Environmental Microbiology 38 (5): 1018–19. PMC 243624. PMID 16345456. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=243624. 
31. ^ Duncan CJG, Pugh N, Pasco DS, Ross SA. (2002). "Isolation of a galactomannan that enhances macrophage activation from the edible fungus Morchella esculenta". Journal of Agricultural and Food Chemistry 50 (20): 5683–85. doi:10.1021/jf020267c. PMID 12236698. 
32. ^ Nitha B, Janardhanan KK. (2008). "Aqueous-ethanolic extract of morel mushroom mycelium Morchella esculenta, protects cisplatin and gentamicin induced nephrotoxicity in mice". Food and Chemical Toxicology 46 (9): 3193–99. doi:10.1016/j.fct.2008.07.007. PMID 18692113. 
33. ^ Rotzoll N, Dunkel A, Hofmann T. (2005). "Activity-guided identification of (S)-malic acid 1-O-D-glucopyranoside (morelid) and gamma-aminobutyric acid as contributors to umami taste and mouth-drying oral sensation of morel mushrooms (Morchella deliciosa Fr.)". Journal of Agricultural and Food Chemistry 53 (10): 4149–56. doi:10.1021/jf050056i. PMID 15884853. 
34. ^ Wasser SP. (2002). "Medicinal mushrooms as a source of antitumor and immunomodulating polysaccharides". Applied Microbiology and Biotechnology 60 (3): 258–74. doi:10.1007/s00253-002-1076-7. PMID 12436306. 
35. ^ Mau JL, Chang CN, Hunag SJ, Chen CC. (2004). "Antioxidant properties of methanolic extracts from Grifola frondosa, Morchella esculenta and Termitomyces albuminosus mycelia". Food Chemistry 87 (1): 111–18. doi:10.1016/j.foodchem.2003.10.026. 
36. ^ Elmastas M, Turkekul I, Ozturk L, Gulcin I, Isildak O, Aboul-Enein HY. (2006). "Antioxidant activity of two wild edible mushrooms (Morchella vulgaris and Morchella esculanta) from North Turkey". Combinatorial Chemistry & High Throughput Screening 9 (6): 443–48. doi:10.2174/138620706777698544. 
37. ^ Gursoy N, Sarikurkcu C, Cengiz M, Solak MH. (2009). "Antioxidant activities, metal contents, total phenolics and flavonoids of seven Morchella species". Food and Chemical Toxicology 47 (9): 2381–88. doi:10.1016/j.fct.2009.06.032. PMID 19563856. 
38. ^ Ying J, Mao X, Ma Q, Zong Y, Wen H. (1987). Icones of Medicinal Fungi from China. Xu Y, Trans.; Science Press: Beijing. pp. 38–45.
39. ^ Zhang G-P, Zhang F, Ru W-M, Han J-R. (2009). "Solid-state fermentation of cornmeal with the ascomycete Morchella esculenta for degrading starch and upgrading nutritional value". World Journal of Microbiology and Biotechnology 26 (1): 15–20. doi:10.1007/s11274-009-0135-y. 
40. ^ Myung K, Narciso JA, Manthey JA. (2008). "Removal of furanocoumarins in grapefruit juice by edible fungi" (PDF). Journal of Agricultural and Food Chemistry 56 (24): 12064–68. doi:10.1021/jf802713g. PMID 19012403. http://ddr.nal.usda.gov/bitstream/10113/22510/1/IND44146736.pdf. Retrieved 2010-03-21. 

External links

• Morchella esculenta in Index Fungorum.
• Morchella esculenta in MycoBank.