Overview

Brief Summary

Grifola Frondosa: Hen of the Woods

Ecology

The Grifola frondosa, or maitake mushroom is a polypore that grows in northern temperate forests, and has a long history of medicinal use. In the western hemisphere, it grows in Canada, and the northeastern and mid-Atlantic US. It is not unheard of, however, to see it in the northwestern or southeastern states.  In the eastern hemisphere, it is found in China, northeastern Japan, and Europe. This mushroom is found on dead or dying trees or stumps, usually around the base. It can also be found growing from submerged, rotting roots. It prefers oaks, though it will also grow on other deciduous hardwoods such as elm, maple, blackgum, and beech (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000). It has also been reported to grow uncommonly on Douglas fir (R. L. Gilbertson, 1986). It tends to prefer the boundary between wooded areas and open fields (Stamets, Mycelium Running, 2005). Maitakes are generally classified as saprophytes, feeding on dead wood. Though they are sometimes found on dying trees, the maitake is rarely the initial pathogen, and will consume already weakened by another organism (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000).

Morphology

            This fungus is a large, soft-fleshed polypore that goes from dark gray-brown when young to lighter gray with age. The fruiting body is a dense cluster of fan-shaped, wavy-margined caps that are 2-10 cm in diameter and extend radially from a common base. Clusters can be anywhere from several to over a hundred pounds in weight. The underside of the caps is covered in 5 mm deep, maze-like pores that extend far down the stem. There are approximately 2 to 4 pores per mm (Kuo, 2010). Hyphal growth is described as fluffy, and non-rhizomorphic, extending in uneven circles when grown on agar. When grown on sawdust or woodchips, hyphae form dense mats that exude orange or rusty fluids (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000). When suspended in liquid culture, growth is a “feather-like mycelial clump” (Bum Chun Lee, 2004). The spores are white, smooth, and elliptical. Cystidia are absent (Kuo, 2010).

History

            Though known and used by many cultures around the world, this mushroom first entered formal Western literature in 1785 under the name Boletus frondosus in James Dickson’s publication Fasciculus Plantarum Cryptogamicarum Britanniae (Harvard University, 2009) . The name was then changed to Grifola frondosa in 1821 by Dickson and fellow British mycologist Samuel Frederick Grey (Index Fungorum Partnership, 2013). The fungus was described in Gray’s textbook, A natural arrangement of British plants as being “eatable, but requir[ing] thorough dressing”, having a “much branched thallus”, and “numerous, halved, smoke grey caps” (Gray, 1821). It currently has many synonyms, including Boletus frondosus, Agaricus frondosus, and Polyporus frondosus (IMA, 2013). Because of its edibility and widely-accepted medicinal uses, the maitake mushroom has many names spanning several languages. It is known as “Laubporling” (or “Ram’s head”) in Germany, the “signorina” (or “lady”) mushroom in Italy, and both as maitake (“dancing mushroom”) and kumotake (“cloud mushroom”) in Japan. In the US it is known as hen-of-the-woods, the dancing butterfly mushroom, or as maitake (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000).

Look-alikes

            The maitake mushroom is often confused with several other species. Its lookalike Grifola umbellata can be distinguished from frondosa by its “multiple caps arising from one stem, a lighter color, and a more fragile texture.” In addition, the spores of G. umbellata are “larger and more cylindrically shaped” (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000). Another misconception is with the mushroom Laetiporus sulphureus because of its similar common name, “chicken-of-the-woods.” The similarity ends there, however, as chicken-of-the-woods is anywhere from bright orange to yellowish-white (Alan E. Bessette, 2007), compared with the light brownish-grey of the maitake.

Cultivation

            Though known by many cultures for many years, efforts have begun only recently to produce it artificially, on a large scale.  Prior to the late 1979, when the first cultivation techniques were developed, maitake mushrooms were available only in the wild (Hobbs, 1996). The first large-scale commercial production began in Japan in 1981, at a modest scale of 325 tons (F. Takama, 1981). For about 10 years, Japan held the monopoly on production, until US and China began producing. Between 1991 and 1997, worldwide production increased around 40-fold, arriving at 331,000 tons produced annually (Chang, 1997).

Medicinal Value

            Traditional medicine systems in countries such as China and Japan have long touted the health benefits of maitake.  One of the earliest mentions of this mushroom is in the Chinese medical text the Shen Nong Ben Cao Jing, from the Han Dynasty (206 BC. – 220 AD.).  Here it is said to be able to “improve the spleen, assuage stomach ailments, treat hemorrhoids, and calm the mind and nerves” (Jones, 1998).  However, modern medicine has only recently begun investigating these claims. While human studies are rare, a single survey of non-human studies showed that studies have been conducted assessing maitake as an anti-diabetic, anti-hypertensive, antineoplastic, antiviral, immunosuppressant, and interferon (Catherine Ulbricht, 2009). A study by the National Cancer Institute agrees with the National Institute of Health of Japan in saying that the powdered fruiting bodies of maitake show significant activity against the HIV virus (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000).

            A study was conducted assessing maitake as a weight-loss aid where mice were either fed a high-cholesterol diet or the same diet supplemented with maitake. After 12 days, the mice given maitake had cholesterol levels 43% lower than those fed the control. By day 24, body weight and body fat were significantly lower as well (K. Kubo, 1996).

            Much of the attention on maitakes is focused on a polysaccharide D-fraction that contains the glucan grifolan (Ohno, 1984). This acid-insoluble, alkali-soluble, hot-water extractable fraction (Konno, 2001) was found to have extreme anti-cancer activity in countless studies. The first was a study where the D-fraction was administered to cancerous mice, and caused complete tumor regression in more than one-third of the trials (Ohno, 1984).  A similar study found that a water extract of maitake inhibited the reproduction of leukemia cells grown in culture (Lovy, 1999). It appears that maitakes have a potential two-fold effect of causing direct apoptosis to cancer cells as well as stimulating the immune system of the host (Konno, 2001). Today, vitamin and supplement companies provide these compounds in the form of “maitake D-fraction” capsules or extracts.

            Another claim often made about maitakes are their ability to reduce blood pressure. A study was conducted where rats were fed either a control diet or one supplemented with 5% maitake powder. The blood pressure of the rats eating the maitakes, after 63 days, was significantly lower than that of the control (Yearul Kabir, 1987).

            Regardless of medical benefits, maitake is widely accepted as a nutritious food. One analysis of 100 grams of dried maitakes contains 377 calories, 26 grams of protein, 19 grams of sugar, and 29 grams of fiber. It is also high in vitamin D, potassium, and niacin (Stamets, Mycelium Running, 2005).

  • Alan E. Bessette, e. a. (2007). Mushrooms of the Southeastern United States. Syracuse: Syracuse University Press.
  • Bum Chun Lee, e. a. (2004). Submerged culture conditions for the production of mycelial biomass and exopolysaccharides by the edible Basidiomycete Grifola frondosa. Enzyme and Microbial Technology, 369-376.
  • Catherine Ulbricht, e. a. (2009). Maitake Mushroom (Grifola frondosa): Systematic Review by the Natural Standard Reseach Collaboration. Journal of the Society for Integrative Oncology, 66-72.
  • Chang, S.-T. (1997). World Production of Cultivated Edible and Medicinal Mushrooms in 1997 with Emphasis on Lentinus edodes (Berk.) Sing, in China. International Journal of Medicinal Mushrooms, 291-300.
  • F. Takama, e. a. (1981). Parenchyma cells, chemical components of maitake mushroom cultured artificially, and their changes by storing and boiling. Mushroom Science, 767-779.
  • Gray, S. F. (1821). A Natural Arrangement of British Plants. London: Baldwin, Cradock, and Joy.
  • Harvard University. (2009, 5 21). Index of Botanical Publications. Retrieved from Harvard University Herbaria: http://kiki.huh.harvard.edu/databases/publication_search.php?mode=details&id=552
  • Hobbs, C. (1996). Medicinal Mushrooms: An Exploration of Tradition, Healing & Culture. Santa Cruz, CA: Botanica Press.
  • IMA. (2013, 9 19). Grifola Frondosa. Retrieved from MycoBank: http://www.mycobank.org/Biolomics.aspx?Table=Mycobank&MycoBankNr_=362177
  • Index Fungorum Partnership. (2013). Grifola frondosa. Retrieved from Index Fungorum: http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=362177
  • Jones, K. (1998). Maitake: A Potential Medicinal Food. Alternative and Complementary Therapies, 420-429.
  • K. Kubo, H. K. (1996). The effect of maitake on liver and serum lipids. Alternative Therapies in Health and Medicine, 62-66.
  • Konno, S. (2001). Maitake D-Fraction: Apoptosis Inducer and Immune Enhancer. Alternative and Complementary Therapies, 102-107.
  • Kuo, M. (2010, March). Grifola Frondosa: The hen of the woods. Retrieved from Mushroom Expert: http://www.mushroomexpert.com/grifola_frondosa.html
  • Lovy, A. (1999). Activity of edible mushrooms against the growth of human T4 leukemia cells. Journal of Herbs, Spices and Medicinal Plants, 49-57.
  • Ohno, N. (1984). Antitumor Activity and Structural Characterization of Glucans Extracted from Cultured Fruit Bodies of Grifola Frondosa. Chemical and Pharmaceutical Bulletin, 1142-1151.
  • R. L. Gilbertson, L. (1986). North American Polypores. Volume 1: Abortiporus - Lindtneria. Journal of Basic Microbiology, 282.
  • Stamets, P. (2000). Growing Gourmet and Medicinal Mushrooms. New York: Crown Publishing.
  • Stamets, P. (2005). Mycelium Running. New York: Ten Speed Press.
  • Yearul Kabir, e. a. (1987). Effect of Shiitake and Maitake Mushrooms on Blood Pressure and Plasma Lipids of Spontaneusly Hypertensive Rats. Journal of Nutritional Science and Vitaminology, 341-346.
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Ecology

Associations

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Broadleaved trees

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Quercus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Castanea sativa

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Corylus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Fagus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Fraxinus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Ilex aquifolium

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Rhododendron ponticum

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Sorbus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Pyrus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Ulmus

Foodplant / parasite
fruitbody of Grifola frondosa parasitises live root of Cedrus

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General Ecology

Ecology

The Grifola frondosa, or maitake mushroom is a polypore that grows in northern temperate forests, and has a long history of medicinal use. In the western hemisphere, it grows in Canada, and the northeastern and mid-Atlantic US. It is not unheard of, however, to see it in the northwestern or southeastern states.  In the eastern hemisphere, it is found in China, northeastern Japan, and Europe. This mushroom is found on dead or dying trees or stumps, usually around the base. It can also be found growing from submerged, rotting roots. It prefers oaks, though it will also grow on other deciduous hardwoods such as elm, maple, blackgum, and beech (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000). It has also been reported to grow uncommonly on Douglas fir (R. L. Gilbertson, 1986). It tends to prefer the boundary between wooded areas and open fields (Stamets, Mycelium Running, 2005). Maitakes are generally classified as saprophytes, feeding on dead wood. Though they are sometimes found on dying trees, the maitake is rarely the initial pathogen, and will consume already weakened by another organism (Stamets, Growing Gourmet and Medicinal Mushrooms, 2000). 

  • Alan E. Bessette, e. a. (2007). Mushrooms of the Southeastern United States. Syracuse: Syracuse University Press.
  • Bum Chun Lee, e. a. (2004). Submerged culture conditions for the production of mycelial biomass and exopolysaccharides by the edible Basidiomycete Grifola frondosa. Enzyme and Microbial Technology, 369-376.
  • Catherine Ulbricht, e. a. (2009). Maitake Mushroom (Grifola frondosa): Systematic Review by the Natural Standard Reseach Collaboration. Journal of the Society for Integrative Oncology, 66-72.
  • Chang, S.-T. (1997). World Production of Cultivated Edible and Medicinal Mushrooms in 1997 with Emphasis on Lentinus edodes (Berk.) Sing, in China. International Journal of Medicinal Mushrooms, 291-300.
  • F. Takama, e. a. (1981). Parenchyma cells, chemical components of maitake mushroom cultured artificially, and their changes by storing and boiling. Mushroom Science, 767-779.
  • Gray, S. F. (1821). A Natural Arrangement of British Plants. London: Baldwin, Cradock, and Joy.
  • Harvard University. (2009, 5 21). Index of Botanical Publications. Retrieved from Harvard University Herbaria: http://kiki.huh.harvard.edu/databases/publication_search.php?mode=details&id=552
  • Hobbs, C. (1996). Medicinal Mushrooms: An Exploration of Tradition, Healing & Culture. Santa Cruz, CA: Botanica Press.
  • IMA. (2013, 9 19). Grifola Frondosa. Retrieved from MycoBank: http://www.mycobank.org/Biolomics.aspx?Table=Mycobank&MycoBankNr_=362177
  • Index Fungorum Partnership. (2013). Grifola frondosa. Retrieved from Index Fungorum: http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=362177
  • Jones, K. (1998). Maitake: A Potential Medicinal Food. Alternative and Complementary Therapies, 420-429.
  • K. Kubo, H. K. (1996). The effect of maitake on liver and serum lipids. Alternative Therapies in Health and Medicine, 62-66.
  • Konno, S. (2001). Maitake D-Fraction: Apoptosis Inducer and Immune Enhancer. Alternative and Complementary Therapies, 102-107.
  • Kuo, M. (2010, March). Grifola Frondosa: The hen of the woods. Retrieved from Mushroom Expert: http://www.mushroomexpert.com/grifola_frondosa.html
  • Lovy, A. (1999). Activity of edible mushrooms against the growth of human T4 leukemia cells. Journal of Herbs, Spices and Medicinal Plants, 49-57.
  • Ohno, N. (1984). Antitumor Activity and Structural Characterization of Glucans Extracted from Cultured Fruit Bodies of Grifola Frondosa. Chemical and Pharmaceutical Bulletin, 1142-1151.
  • R. L. Gilbertson, L. (1986). North American Polypores. Volume 1: Abortiporus - Lindtneria. Journal of Basic Microbiology, 282.
  • Stamets, P. (2000). Growing Gourmet and Medicinal Mushrooms. New York: Crown Publishing.
  • Stamets, P. (2005). Mycelium Running. New York: Ten Speed Press.
  • Yearul Kabir, e. a. (1987). Effect of Shiitake and Maitake Mushrooms on Blood Pressure and Plasma Lipids of Spontaneusly Hypertensive Rats. Journal of Nutritional Science and Vitaminology, 341-346.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage: Grifola frondosa

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Specimens with Barcodes: 1
Species With Barcodes: 1
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Wikipedia

Grifola frondosa

"Ram's Head" redirects here. For the mountain, see Rams Head. For the military skill badge, see Ram's Head Device. For the head of the male animal, see sheep.

Grifola frondosa is a polypore mushroom that grows in clusters at the base of trees, particularly oaks. The mushroom is commonly known among English speakers as hen-of-the-woods, ram's head and sheep's head. In the United States' supplement market, as well as in Asian grocery stores, the mushroom is known by its Japanese name maitake (舞茸), which means "dancing mushroom". Throughout Italian American communities in the northeastern United States, it is commonly known as the signorina mushroom. G. frondosa should not be confused with Laetiporus sulphureus, another edible bracket fungus that is commonly called chicken of the woods or "sulphur shelf". The fungus becomes inedible like all polypores when they are older, because it is too tough to eat.

The fungus is native to the northeastern part of Japan and North America, and is prized in traditional Chinese and Japanese herbology as a medicinal mushroom, an aid to balance out altered body systems to a normal level. It is widely eaten in Japan, and its popularity in western cuisine is growing, although the mushroom has been alleged to cause allergic reactions in rare cases.

Grifola frondosa
View the Mycomorphbox template that generates the following list
Mycological characteristics
pores on hymenium

cap is offset

or indistinct
hymenium is decurrent
lacks a stipe
spore print is white
ecology is parasitic
edibility: choice

Description[edit]

Like the sulphur shelf mushroom, G. frondosa is a perennial fungus that often grows in the same place for a number of years in succession. It occurs most prolifically in the northeastern regions of the United States, but has been found as far west as Idaho.

G. frondosa grows from an underground tuber-like structure known as a sclerotium, about the size of a potato. The fruiting body, occurring as large as 100 cm, is a cluster consisting of multiple grayish-brown caps which are often curled or spoon-shaped, with wavy margins and 2–7 cm broad. The undersurface of each cap bears approximately one to three pores per millimeter, with the tubes rarely deeper than 3 mm. The milky-white stipe (stalk) has a branchy structure and becomes tough as the mushroom matures.

In Japan, the Maitake can grow to more than 100 pounds (40 kilograms), earning this giant mushroom the title "King of Mushrooms". Maitake is one of the major culinary mushrooms used in Japan, the others being shiitake, shimeji and enoki. They are used in a wide variety of dishes, often being a key ingredient in nabemono or cooked in foil with butter.

Use in traditional Eastern medicine[edit]

The sclerotia from which hen of the woods arises have been used in traditional Chinese and Japanese medicine to enhance the immune system. Researchers have also indicated that whole maitake has the ability to regulate blood pressure, glucose, insulin, and both serum and liver lipids, such as cholesterol, triglycerides, and phospholipids, and may also be useful for weight loss.[citation needed]

Maitake is rich in minerals (such as potassium, calcium, and magnesium), various vitamins (B2, D2 and niacin), fibers and amino acids. One active constituent in Maitake for enhancing the immune activity was identified in the late 1980s as a protein-bound beta-glucan compound.

Maitake research[edit]

In 2009, a phase I/II human trial, conducted by Memorial Sloan–Kettering Cancer Center, showed Maitake could stimulate the immune systems of breast cancer patients.[2] Small experiments with human cancer patients have shown Maitake can stimulate immune system cells, like NK cells.[3][4] In vitro research has also shown Maitake can stimulate immune system cells.[5] An in vivo experiment showed that Maitake could stimulate both the innate immune system and adaptive immune system.[6]

In vitro research has shown Maitake can induce apoptosis in various cancer cell lines as well as inhibit the growth of various types of cancer cells.[7] Small studies with human cancer patients revealed that a portion of the Maitake mushroom, known as the "Maitake D-fraction", possesses anti-cancer activity.[8][9] In vitro research demonstrated the mushroom has potential anti-metastatic properties.[10]

Research has shown Maitake has a hypoglycemic effect, and may be beneficial for the management of diabetes.[7] The reason Maitake lowers blood sugar is because the mushroom naturally contains an alpha glucosidase inhibitor.[11]

Maitake contains antioxidants and may partially inhibit the enzyme cyclooxygenase.[12] An experiment showed that an extract of Maitake inhibited angiogenesis via inhibition of the vascular endothelial growth factor (VEGF).[13]

Lys-N is a unique protease found in Maitake.[14] Lys-N is used for proteomics experiments due to its protein cleavage specificity.[15]

See also[edit]

References[edit]

  1. ^ McIlvaine, Charles; Robert K. Macadam; and Robert L. Shaffer. 1973. One Thousand American Fungi. Dover Publications. New York. 729 pp. (Polyporus frondosus, pp. 482-483 & Plate CXXVIII.)
  2. ^ Deng G, Lin H, Seidman A, et al. (September 2009). "A phase I/II trial of a polysaccharide extract from Grifola frondosa (Maitake mushroom) in breast cancer patients: immunological effects". Journal of Cancer Research and Clinical Oncology 135 (9): 1215–21. doi:10.1007/s00432-009-0562-z. PMID 19253021. 
  3. ^ Kodama N, Komuta K, Nanba H (2003). "Effect of Maitake (Grifola frondosa) D-Fraction on the activation of NK cells in cancer patients". Journal of Medicinal Food 6 (4): 371–7. doi:10.1089/109662003772519949. PMID 14977447. 
  4. ^ Kodama N, Komuta K, Sakai N, Nanba H (December 2002). "Effects of D-Fraction, a polysaccharide from Grifola frondosa on tumor growth involve activation of NK cells". Biological & Pharmaceutical Bulletin 25 (12): 1647–50. doi:10.1248/bpb.25.1647. PMID 12499658. 
  5. ^ Kodama N, Asakawa A, Inui A, Masuda Y, Nanba H (March 2005). "Enhancement of cytotoxicity of NK cells by D-Fraction, a polysaccharide from Grifola frondosa". Oncology Reports 13 (3): 497–502. doi:10.3892/or.13.3.497. PMID 15706424. 
  6. ^ Kodama N, Murata Y, Nanba H (2004). "Administration of a polysaccharide from Grifola frondosa stimulates immune function of normal mice". Journal of Medicinal Food 7 (2): 141–5. doi:10.1089/1096620041224012. PMID 15298759. 
  7. ^ a b Ulbricht C, Weissner W, Basch E, Giese N, Hammerness P, Rusie-Seamon E, Varghese M, Woods J. (2009). "Maitake mushroom (Grifola frondosa): systematic review by the natural standard research collaboration". Journal of the Society for Integrative Oncology 7 (2): 66–72. PMID 19476741. 
  8. ^ Kodama N, Komuta K, Nanba H (June 2002). "Can maitake MD-fraction aid cancer patients?". Alternative Medicine Review 7 (3): 236–9. PMID 12126464. 
  9. ^ Nanba H, Kubo K (December 1997). "Effect of Maitake D-fraction on cancer prevention". Annals of the New York Academy of Sciences 833 (1 Cancer): 204–7. doi:10.1111/j.1749-6632.1997.tb48611.x. PMID 9616756. 
  10. ^ Masuda Y, Murata Y, Hayashi M, Nanba H (June 2008). "Inhibitory effect of MD-Fraction on tumor metastasis: involvement of NK cell activation and suppression of intercellular adhesion molecule (ICAM)-1 expression in lung vascular endothelial cells". Biological & Pharmaceutical Bulletin 31 (6): 1104–8. doi:10.1248/bpb.31.1104. PMID 18520039. 
  11. ^ Matsuur H, Asakawa C, Kurimoto M, Mizutani J (July 2002). "Alpha-glucosidase inhibitor from the seeds of balsam pear (Momordica charantia) and the fruit bodies of Grifola frondosa". Bioscience, Biotechnology, and Biochemistry 66 (7): 1576–8. doi:10.1271/bbb.66.1576. PMID 12224646. 
  12. ^ Zhang Y, Mills GL, Nair MG (December 2002). "Cyclooxygenase inhibitory and antioxidant compounds from the mycelia of the edible mushroom Grifola frondosa". Journal of Agricultural and Food Chemistry 50 (26): 7581–5. doi:10.1021/jf0257648. PMID 12475274. 
  13. ^ Lee JS, Park BC, Ko YJ, et al. (December 2008). "Grifola frondosa (maitake mushroom) water extract inhibits vascular endothelial growth factor-induced angiogenesis through inhibition of reactive oxygen species and extracellular signal-regulated kinase phosphorylation". Journal of Medicinal Food 11 (4): 643–51. doi:10.1089/jmf.2007.0629. PMID 19053855. 
  14. ^ Nonaka, T; Y Hashimoto; K Takio (July 1998). "Kinetic characterization of lysine-specific metalloendopeptidases from Grifola frondosa and Pleurotus ostreatus fruiting bodies". Journal of Biochemistry 124 (1): 157–162. doi:10.1093/oxfordjournals.jbchem.a022074. ISSN 0021-924X. PMID 9644258. 
  15. ^ Taouatas, Nadia; Madalina M Drugan; Albert J R Heck; Shabaz Mohammed (May 2008). "Straightforward ladder sequencing of peptides using a Lys-N metalloendopeptidase". Nat Meth 5 (5): 405–407. doi:10.1038/nmeth.1204. ISSN 1548-7091. PMID 18425140. 
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