Polyozellus
Encyclopedia
Polyozellus is a fungal genus
in the family
Thelephoraceae, a grouping of mushroom
s known collectively as the leathery earthfans. A monotypic genus, it contains the single species Polyozellus multiplex, first described in 1899, and commonly
known as the blue chanterelle, the clustered blue chanterelle, or, in Alaska
, the black chanterelle. The distinctive fruit body
of this species comprises blue- to purple-colored clusters of vase- or spoon-shaped caps
with veiny wrinkles on the undersurface that run down the length of the stem
.
Polyozellus has had a varied taxonomic
history and has been reclassified several times at both the family
and genus level. The range
of Polyozellus includes North America and eastern Asia, where P. multiplex may be found growing on the ground in coniferous forests, usually under spruce
and fir
trees. It is an edible
species, and has been harvested for commercial purposes. Polyozellus multiplex contains the bioactive
compound polyozellin
, shown to have various physiological
properties, including suppressive effects on stomach cancer
.
of the species was written by botanist Lucien M. Underwood
in 1899, based on a specimen found the previous year in the woods of Mount Desert, Maine
. Although he called the new species a Cantharellus
, he noted that "the plant is a remarkable one and from its habit might well form a distinct genus since it has little in common with Cantharellus except its fold-like gills." In 1910, William Murrill
transferred it to the new genus Polyozellus; Murrill thought the compound structure of the stem to be a sufficiently unique characteristic to warrant it being separated from Cantharellus species, which have simpler stem structures. In 1920, specimens from a Japanese collection compiled by A. Yasuda were sent to mycologist Curtis Gates Lloyd
, who believed the fungus to be a new species and named it Phyllocarbon yasudai.
No further collections of the fungus were reported until 1937, when it was found in Quebec
, Canada. The next year, Paul Shope considered the genus Polyozellus to be superfluous, pointed out that the compound fruit bodies
and the wrinkled hymenium
were instead consistent with the genus Craterellus
. In 1939, American mycologist Lee Overholts, in a letter to the journal Mycologia
, opined that both of these authors had overlooked a 1925 publication by Calvin Kauffman, who made notes and photos of the species collected in the Rocky Mountains
of Wyoming
and Colorado
, and in the Cascade Mountains
of Washington and Oregon
. Kauffman believed the species to be merely "a very extreme growth condition" of Cantharellus clavatus (now known as Gomphus clavatus
) and suggested there was no reason for transferring the species to the genus Craterellus. Mycologists Alexander H. Smith and Elizabeth Morse, in their 1947 publication on Cantharellus species in the United States, placed the species in a new section Polyozellus, but did not separate it from the genus Cantharellus; they defined the distinguishing characteristics of Polyozellus as the small, roughened, hyaline
spores and the color change of the flesh in potassium hydroxide
solution, adding that "the spores are unusual for the genus but in our estimation do not warrant excluding the species."
In 1953, Rokuya Imazeki took into consideration differences in spore
characteristics: species in the genus Cantharellus were not known to have spores that were subglobose (roughly spherical) and tuberculate (covered with wart-like projections) like Polyozellus; however, these spore characteristics were common in species in the Thelephoraceae family (Cantharellus belongs in a different family, the Cantharellaceae
). Other characteristics linking the blue chanterelle with the Thelephoraceae included the dark color, the strong odor (especially in dried specimens), and the presence of thelephoric acid
, a mushroom pigment
common in the family. Taken together, these factors led Imazeki to propose the new family Phylacteriaceae. The suggested family-level taxonomical
change was not accepted by other authors; for example, in 1954, Seiichi Kawamura renamed it Thelephora multiplex. As of 2009, both Index Fungorum
and MycoBank
list Polyozellus as being within the Thelephoraceae family, a grouping of mushrooms commonly known as the leathery earthfans. The genus name is derived from the Greek
poly meaning many, and oz, meaning branch. The specific epithet multiplex means "in many pieces", referring to the compound nature of the fruit body. Common name
s for this species include the blue chanterelle and the clustered blue chanterelle. In Alaska
, where specimens typically have very dark-colored fruit bodies, it is called the black chanterelle, although this name is shared with some Craterellus
species.
Polyozellus multiplex is part of the group of fungi collectively known as cantharelloid mushrooms (which includes the genera Cantharellus
, Craterellus, Gomphus
, and Polyozellus) because of the similarity of their fruit body structures and the morphology
of the spore-producing region (the hymenophore
) on the underside of the caps. The fan- or funnel-shaped fruit bodies of the black chanterelle grow clustered together on the ground, often in large masses that may reach aggregate diameters of up to 1 metres (3.3 ft), although they are usually up to 30 centimetres (11.8 in).
The individual caps
, 3 – wide and almost as long, are violet-black, with edges that are initially whitish, and with a glaucous
surface—a white powdery accumulation of spore deposit. The upper surface may be zonate—lined with what appear to be multiple concentric zones of texture caused by areas of fine hairs (a tomentum
); and the edges of the caps have a layer of very fine hairs and are lobed and wavy. The underside of the caps bears the fertile, spore-making tissue called the hymenium
, which typically has shallow, crowded wrinkles or veins that are roughly the same color or paler than the top surface. Some variation in color has been observed depending on the collection location. For example, specimens found in Alaska are more likely to be jet-black in color with a dark gray underside.
Fruit bodies may be up to 15 cm (5.9 in) high (including the stem) and 10 cm (3.9 in) wide. Occasionally, much larger clusters of fused mushrooms are found, up to a meter in diameter. The stem
is dark purplish-black with a smooth (glabrous
) and dry surface; the stems are often fused at the base. It is typically 1.5 – wide and up to 5 cm (2 in) long. The flesh
is dark violet, soft but breaking easily. The spore deposit
is white.
The spores
are roughly spherical to broadly ellipsoid in shape, covered with small wart-like projections (tubercules), and have dimensions of 6–8.5 by 5.5–8 µm
. Viewed microscopically, they are hyaline
, meaning they appear translucent or colorless. Chemical tests may also be used to help distinguish the spores: in the presence of potassium hydroxide
(KOH), the spores turn slightly green; the spores are not amyloid
, meaning they do not take iodine
when treated with Melzer's reagent
; the spores are acyanophilous, meaning they do not readily absorb methyl blue
stain. The cystidia
that comprise the hymenium
are filamentous and 3–4 µm wide by 28–40 µm long. The outer tissue layer of the cap—the cuticle, or pileipellis
—is made of interwoven hypha
e, and stains olive-green in KOH. Clamp connection
s are present, but not at all the cell partitions
. The basidia
, the spore-bearing cells, are 32–38 by 5–6 µm and four-spored.
) also has a blackish fruit body and a smooth hymenium, but is distinguished from P. multiplex by its thin flesh, a trumpet- or tubular-shaped fruit body (rather than fan- or spoon-shaped), and grey to black colors. A closely related species, the fragrant chanterelle (Cantharellus odoratus), also tends to grow in dense clusters, but it is orange rather than blue. Craterellus caeruleofuscus
does not form compound clusters, and is not restricted to coniferous forests. The pig's ear Gomphus, species Gomphus clavatus
, is similar in shape and form but fleshier, and light violet to pink.
e of the fungus grow in a mutualistic association with the roots of plants, but the fungal hyphae generally do not penetrate the cells of the plant's roots. The species grows in coniferous woods under spruce
and fir
, and more frequently at higher elevations. It is most often encountered in summer and fall.
This species is northern and alpine
in distribution, and rarely encountered. Collections have been made in the United States (including Maine
, Oregon
, Colorado
, New Mexico
, and Alaska), Canada (Quebec
and British Columbia
), China, Japan, and Korea. The disjunct distribution
of this species in North America and East Asia has been noted to occur in a number of other fungal species as well. Polyozellus multiplex is also found in the Queen Charlotte Islands
, where it is commercially harvested.
, and is collected for sale in Asian countries such as Korea, Japan, and China. In North America, it is sometimes collected recreationally, and commercially. The taste is described as mild, and the odor as mild or aromatic. Mycologist David Arora
claims the flavor to be inferior to Craterellus. Fruit bodies may be preserved by drying.
The compound polyozellin
—a chemical which can be isolated and purified from P. multiplex—inhibits prolyl endopeptidase
(PEP), an enzyme
that has a role in processing proteins (specifically, amyloid precursor protein
) in Alzheimer's disease
. Chemicals that inhibit PEP have attracted research interest due to their potential therapeutic effects. Further analyses of extracts from P. multiplex revealed similar dibenzofuran
yl derivatives
of polyozellin, each with different chemical properties
, including kynapcin
-12, kynapcin-13 and -28, and -24. A total synthesis
of kynapcin-24 was achieved in 2009.
. The study showed that feeding a low concentration (0.5% or 1%) of the mushroom extract enhanced the activities of the enzymes glutathione S-transferase
and superoxide dismutase
, and increased the abundance of the molecule glutathione
. The extract also augmented the expression of the protein p53
. All of these substances protect the human organism against cancer. Additional studies reported in 2004 and 2006 attribute anti-tumor properties to polyozellin.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Thelephoraceae, a grouping of mushroom
Mushroom
A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or on its food source. 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 that...
s known collectively as the leathery earthfans. A monotypic genus, it contains the single species Polyozellus multiplex, first described in 1899, and commonly
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
known as the blue chanterelle, the clustered blue chanterelle, or, in Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
, the black chanterelle. The distinctive fruit body
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
of this species comprises blue- to purple-colored clusters of vase- or spoon-shaped caps
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
with veiny wrinkles on the undersurface that run down the length of the stem
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
.
Polyozellus has had a varied taxonomic
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
history and has been reclassified several times at both the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
and genus level. The range
Range (biology)
In biology, the range or distribution of a species is the geographical area within which that species can be found. Within that range, dispersion is variation in local density.The term is often qualified:...
of Polyozellus includes North America and eastern Asia, where P. multiplex may be found growing on the ground in coniferous forests, usually under spruce
Spruce
A spruce is a tree of the genus Picea , a genus of about 35 species of coniferous evergreen trees in the Family Pinaceae, found in the northern temperate and boreal regions of the earth. Spruces are large trees, from tall when mature, and can be distinguished by their whorled branches and conical...
and fir
Fir
Firs are a genus of 48–55 species of evergreen conifers in the family Pinaceae. They are found through much of North and Central America, Europe, Asia, and North Africa, occurring in mountains over most of the range...
trees. It is an edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...
species, and has been harvested for commercial purposes. Polyozellus multiplex contains the bioactive
Biological activity
In pharmacology, biological activity or pharmacological activity describes the beneficial or adverse effects of a drug on living matter. When a drug is a complex chemical mixture, this activity is exerted by the substance's active ingredient or pharmacophore but can be modified by the other...
compound polyozellin
Polyozellin
Polyozellin is a chemical which occurs in the mushroom Polyozellus multiplex. It inhibits prolyl endopeptidase , an enzyme that has a role in processing proteins in Alzheimer's disease. Chemicals that inhibit prolyl endopeptidase have attracted research interest due to their potential therapeutic...
, shown to have various physiological
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...
properties, including suppressive effects on stomach cancer
Stomach cancer
Gastric cancer, commonly referred to as stomach cancer, can develop in any part of the stomach and may spread throughout the stomach and to other organs; particularly the esophagus, lungs, lymph nodes, and the liver...
.
History and taxonomy
The first published descriptionSpecies description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...
of the species was written by botanist Lucien M. Underwood
Lucien Marcus Underwood
Lucien Marcus Underwood was an American botanist and mycologist of the 19th and early 20th centuries.-Biography:He was born in New Woodstock, New York, and graduated from Syracuse University...
in 1899, based on a specimen found the previous year in the woods of Mount Desert, Maine
Mount Desert, Maine
Mount Desert is a town on Mount Desert Island in Hancock County, Maine, United States. The population was 2,109 at the 2000 census. Incorporated in 1789, the town currently encompasses the villages of Otter Creek, Seal Harbor, Northeast Harbor, Somesville, Hall Quarry, and Pretty...
. Although he called the new species a Cantharellus
Cantharellus
Cantharellus is a genus of popular edible mushrooms, commonly known as chanterelles . They are mycorrhizal fungi, meaning they form symbiotic associations with plants, making them very difficult to cultivate...
, he noted that "the plant is a remarkable one and from its habit might well form a distinct genus since it has little in common with Cantharellus except its fold-like gills." In 1910, William Murrill
William Murrill
William Alphonso Murrill was an American mycologist, known for his contributions to the knowledge of the Agaricales and Polyporaceae.- Education :...
transferred it to the new genus Polyozellus; Murrill thought the compound structure of the stem to be a sufficiently unique characteristic to warrant it being separated from Cantharellus species, which have simpler stem structures. In 1920, specimens from a Japanese collection compiled by A. Yasuda were sent to mycologist Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...
, who believed the fungus to be a new species and named it Phyllocarbon yasudai.
Quebec
Quebec or is a province in east-central Canada. It is the only Canadian province with a predominantly French-speaking population and the only one whose sole official language is French at the provincial level....
, Canada. The next year, Paul Shope considered the genus Polyozellus to be superfluous, pointed out that the compound fruit bodies
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
and the wrinkled hymenium
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 others some cells develop into sterile cells called cystidia or...
were instead consistent with the genus Craterellus
Craterellus
Craterellus is a genus of generally edible fungi similar to the closely related chanterelles, with some species recently reassigned to this genus. They are distinguished by their lack of gill-like structures on the underside of their caps....
. In 1939, American mycologist Lee Overholts, in a letter to the journal Mycologia
Mycologia
Mycologia is a peer-reviewed scientific journal that publishes papers on all aspects of the fungi, including lichens. It first appeared as a bimonthly journal in January of 1909, published by the New York Botanical Garden under the editorship of William Murrill. It became the official journal of...
, opined that both of these authors had overlooked a 1925 publication by Calvin Kauffman, who made notes and photos of the species collected in the Rocky Mountains
Rocky Mountains
The Rocky Mountains are a major mountain range in western North America. The Rocky Mountains stretch more than from the northernmost part of British Columbia, in western Canada, to New Mexico, in the southwestern United States...
of Wyoming
Wyoming
Wyoming is a state in the mountain region of the Western United States. The western two thirds of the state is covered mostly with the mountain ranges and rangelands in the foothills of the Eastern Rocky Mountains, while the eastern third of the state is high elevation prairie known as the High...
and Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...
, and in the Cascade Mountains
Cascade Range
The Cascade Range is a major mountain range of western North America, extending from southern British Columbia through Washington and Oregon to Northern California. It includes both non-volcanic mountains, such as the North Cascades, and the notable volcanoes known as the High Cascades...
of Washington and Oregon
Oregon
Oregon is a state in the Pacific Northwest region of the United States. It is located on the Pacific coast, with Washington to the north, California to the south, Nevada on the southeast and Idaho to the east. The Columbia and Snake rivers delineate much of Oregon's northern and eastern...
. Kauffman believed the species to be merely "a very extreme growth condition" of Cantharellus clavatus (now known as Gomphus clavatus
Gomphus clavatus
Gomphus clavatus is an edible species of fungus in the genus Gomphus, family Gomphaceae. It is commonly known as pig's ears, alluding to the violet underside and yellowish cap of the fruit bodies. Other common names for this species include clustered chanterelle and violet chanterelle...
) and suggested there was no reason for transferring the species to the genus Craterellus. Mycologists Alexander H. Smith and Elizabeth Morse, in their 1947 publication on Cantharellus species in the United States, placed the species in a new section Polyozellus, but did not separate it from the genus Cantharellus; they defined the distinguishing characteristics of Polyozellus as the small, roughened, hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
spores and the color change of the flesh in potassium hydroxide
Potassium hydroxide
Potassium hydroxide is an inorganic compound with the formula KOH, commonly called caustic potash.Along with sodium hydroxide , this colorless solid is a prototypical strong base. It has many industrial and niche applications. Most applications exploit its reactivity toward acids and its corrosive...
solution, adding that "the spores are unusual for the genus but in our estimation do not warrant excluding the species."
In 1953, Rokuya Imazeki took into consideration differences in spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...
characteristics: species in the genus Cantharellus were not known to have spores that were subglobose (roughly spherical) and tuberculate (covered with wart-like projections) like Polyozellus; however, these spore characteristics were common in species in the Thelephoraceae family (Cantharellus belongs in a different family, the Cantharellaceae
Cantharellaceae
The Cantharellaceae are a family of fungi in the order Cantharellales. The family contains the chanterelles and related species, a group of fungi that superficially resemble agarics but have smooth, wrinkled, or gill-like hymenophores...
). Other characteristics linking the blue chanterelle with the Thelephoraceae included the dark color, the strong odor (especially in dried specimens), and the presence of thelephoric acid
Thelephoric acid
right|thumb|Thelephoric acidThelephoric acid is a terphenylquinone pigment which is found in several fungi, such as Omphalotus subilludens and Polyozellus multiplex. Thelephoric acid has been shown to inhibit prolyl endopeptidase, an enzyme that has a role in processing proteins in Alzheimer's...
, a mushroom pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...
common in the family. Taken together, these factors led Imazeki to propose the new family Phylacteriaceae. The suggested family-level taxonomical
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
change was not accepted by other authors; for example, in 1954, Seiichi Kawamura renamed it Thelephora multiplex. As of 2009, both Index Fungorum
Index Fungorum
Index Fungorum, an international project to index all formal names in the Fungi Kingdom. Somewhat comparable to the IPNI, but with more contributing institutions....
and MycoBank
MycoBank
MycoBank is an online database, documenting new mycological names and combinations, eventually combined with descriptions and illustrations. It is run by the Centraalbureau voor Schimmelcultures fungal biodiversity center in Utrecht....
list Polyozellus as being within the Thelephoraceae family, a grouping of mushrooms commonly known as the leathery earthfans. The genus name is derived from the Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
poly meaning many, and oz, meaning branch. The specific epithet multiplex means "in many pieces", referring to the compound nature of the fruit body. Common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...
s for this species include the blue chanterelle and the clustered blue chanterelle. In Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
, where specimens typically have very dark-colored fruit bodies, it is called the black chanterelle, although this name is shared with some Craterellus
Craterellus
Craterellus is a genus of generally edible fungi similar to the closely related chanterelles, with some species recently reassigned to this genus. They are distinguished by their lack of gill-like structures on the underside of their caps....
species.
Description
Cantharellus
Cantharellus is a genus of popular edible mushrooms, commonly known as chanterelles . They are mycorrhizal fungi, meaning they form symbiotic associations with plants, making them very difficult to cultivate...
, Craterellus, Gomphus
Gomphus (fungus)
Gomphus is a small genus of cantharelloid fungi in the family Gomphaceae. The genus has a widespread distribution in temperate regions, and contains 10 species. Once presumed to be related to chanterelles, molecular study has shown them to be allied with stinkhorns and fairy clubs...
, and Polyozellus) because of the similarity of their fruit body structures and the morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
of the spore-producing region (the hymenophore
Hymenophore
A hymenophore refers to the hymenium-bearing structure of a fungal fruiting body. Hymenophores can be smooth surfaces, lamellae, folds, tubes, or teeth....
) on the underside of the caps. The fan- or funnel-shaped fruit bodies of the black chanterelle grow clustered together on the ground, often in large masses that may reach aggregate diameters of up to 1 metres (3.3 ft), although they are usually up to 30 centimetres (11.8 in).
The individual caps
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
, 3 – wide and almost as long, are violet-black, with edges that are initially whitish, and with a glaucous
Glaucous
Glaucous is used to describe the pale grey or bluish-green appearance of the surfaces of some plants, as well as in the names of birds, such as the Glaucous Gull , Glaucous-winged Gull , Glaucous Macaw , and Glaucous...
surface—a white powdery accumulation of spore deposit. The upper surface may be zonate—lined with what appear to be multiple concentric zones of texture caused by areas of fine hairs (a tomentum
Tomentum
Tomentum may refer to the following:*In botany, a covering of closely matted or fine hairs on plant leaves. *A network of minute blood vessels in the brain.* Tomentum in zoology are a short, soft pubescence...
); and the edges of the caps have a layer of very fine hairs and are lobed and wavy. The underside of the caps bears the fertile, spore-making tissue called the hymenium
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 others some cells develop into sterile cells called cystidia or...
, which typically has shallow, crowded wrinkles or veins that are roughly the same color or paler than the top surface. Some variation in color has been observed depending on the collection location. For example, specimens found in Alaska are more likely to be jet-black in color with a dark gray underside.
Fruit bodies may be up to 15 cm (5.9 in) high (including the stem) and 10 cm (3.9 in) wide. Occasionally, much larger clusters of fused mushrooms are found, up to a meter in diameter. The stem
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
is dark purplish-black with a smooth (glabrous
Glabrousness
Glabrousness is the technical term for an anatomically atypical lack of hair, down, or similar structures...
) and dry surface; the stems are often fused at the base. It is typically 1.5 – wide and up to 5 cm (2 in) long. The flesh
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....
is dark violet, soft but breaking easily. The spore deposit
Spore print
thumb|300px|right|Making a spore print of the mushroom Volvariella volvacea shown in composite: mushroom cap laid on white and dark paper; cap removed after 24 hours showing pinkish-tan spore print...
is white.
Microscopic characteristics
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...
are roughly spherical to broadly ellipsoid in shape, covered with small wart-like projections (tubercules), and have dimensions of 6–8.5 by 5.5–8 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
. Viewed microscopically, they are hyaline
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...
, meaning they appear translucent or colorless. Chemical tests may also be used to help distinguish the spores: in the presence of potassium hydroxide
Potassium hydroxide
Potassium hydroxide is an inorganic compound with the formula KOH, commonly called caustic potash.Along with sodium hydroxide , this colorless solid is a prototypical strong base. It has many industrial and niche applications. Most applications exploit its reactivity toward acids and its corrosive...
(KOH), the spores turn slightly green; the spores are not amyloid
Amyloid
Amyloids are insoluble fibrous protein aggregates sharing specific structural traits. Abnormal accumulation of amyloid in organs may lead to amyloidosis, and may play a role in various neurodegenerative diseases.-Definition:...
, meaning they do not take iodine
Iodine
Iodine is a chemical element with the symbol I and atomic number 53. The name is pronounced , , or . The name is from the , meaning violet or purple, due to the color of elemental iodine vapor....
when treated with Melzer's reagent
Melzer's Reagent
Melzer's reagent is a chemical reagent used by mycologists to assist with the identification of fungi.-Composition:...
; the spores are acyanophilous, meaning they do not readily absorb methyl blue
Methyl blue
Methyl blue, also known as Cotton blue, Helvetia blue, Acid blue 93, or C.I. 42780, is a chemical compound with the molecular formula C37H27N3Na2O9S3. It is used as a stain in histology. Methyl blue stains collagen blue in tissue sections. It is soluble in water and slightly soluble in ethanol...
stain. The cystidia
Cystidium
A cystidium is a relatively large cell found on the hymenium of a basidiomycete , often between clusters of basidia. Since cystidia have highly varied and distinct shapes that are often unique to a particular species or genus, they are a useful micromorphological characteristic in the...
that comprise the hymenium
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 others some cells develop into sterile cells called cystidia or...
are filamentous and 3–4 µm wide by 28–40 µm long. The outer tissue layer of the cap—the cuticle, or pileipellis
Pileipellis
thumb|300px||right|The cuticle of some mushrooms, such as [[Russula mustelina]] shown here, can be peeled from the cap, and may be useful as an identification feature....
—is made of interwoven hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e, and stains olive-green in KOH. Clamp connection
Clamp connection
A clamp connection is a structure formed by growing hyphal cells of certain fungi. It is created to ensure each septum, or segment of hypha separated by crossed walls, receives a set of differing nuclei, which are obtained through mating of hyphae of differing sexual types...
s are present, but not at all the cell partitions
Septum
In anatomy, a septum is a wall, dividing a cavity or structure into smaller ones.-In human anatomy:...
. The basidia
Basidium
thumb|right|500px|Schematic showing a basidiomycete mushroom, gill structure, and spore-bearing basidia on the gill margins.A basidium is a microscopic, spore-producing structure found on the hymenophore of fruiting bodies of basidiomycete fungi. The presence of basidia is one of the main...
, the spore-bearing cells, are 32–38 by 5–6 µm and four-spored.
Similar species
The horn-of-plenty mushroom (Craterellus cornucopioidesCraterellus cornucopioides
Craterellus cornucopioides is an edible mushroom, also known as trumpet of death, black chanterelle, black trumpet, or horn of plenty. The Cornucopia, in Greek mythology, referred to the magnificent horn of the goat Amalthea, that filled itself with whatever meat or drink its owner requested...
) also has a blackish fruit body and a smooth hymenium, but is distinguished from P. multiplex by its thin flesh, a trumpet- or tubular-shaped fruit body (rather than fan- or spoon-shaped), and grey to black colors. A closely related species, the fragrant chanterelle (Cantharellus odoratus), also tends to grow in dense clusters, but it is orange rather than blue. Craterellus caeruleofuscus
Craterellus caeruleofuscus
Craterellus caeruleofuscus is a species of fungus in the Cantharellaceae family of mushrooms.-External links:*...
does not form compound clusters, and is not restricted to coniferous forests. The pig's ear Gomphus, species Gomphus clavatus
Gomphus clavatus
Gomphus clavatus is an edible species of fungus in the genus Gomphus, family Gomphaceae. It is commonly known as pig's ears, alluding to the violet underside and yellowish cap of the fruit bodies. Other common names for this species include clustered chanterelle and violet chanterelle...
, is similar in shape and form but fleshier, and light violet to pink.
Habitat and distribution
Polyozellus multiplex is an ectomycorrhizal species, meaning that the hyphaHypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...
e of the fungus grow in a mutualistic association with the roots of plants, but the fungal hyphae generally do not penetrate the cells of the plant's roots. The species grows in coniferous woods under spruce
Spruce
A spruce is a tree of the genus Picea , a genus of about 35 species of coniferous evergreen trees in the Family Pinaceae, found in the northern temperate and boreal regions of the earth. Spruces are large trees, from tall when mature, and can be distinguished by their whorled branches and conical...
and fir
Fir
Firs are a genus of 48–55 species of evergreen conifers in the family Pinaceae. They are found through much of North and Central America, Europe, Asia, and North Africa, occurring in mountains over most of the range...
, and more frequently at higher elevations. It is most often encountered in summer and fall.
This species is northern and alpine
Alpine climate
Alpine climate is the average weather for a region above the tree line. This climate is also referred to as mountain climate or highland climate....
in distribution, and rarely encountered. Collections have been made in the United States (including Maine
Maine
Maine is a state in the New England region of the northeastern United States, bordered by the Atlantic Ocean to the east and south, New Hampshire to the west, and the Canadian provinces of Quebec to the northwest and New Brunswick to the northeast. Maine is both the northernmost and easternmost...
, Oregon
Oregon
Oregon is a state in the Pacific Northwest region of the United States. It is located on the Pacific coast, with Washington to the north, California to the south, Nevada on the southeast and Idaho to the east. The Columbia and Snake rivers delineate much of Oregon's northern and eastern...
, Colorado
Colorado
Colorado is a U.S. state that encompasses much of the Rocky Mountains as well as the northeastern portion of the Colorado Plateau and the western edge of the Great Plains...
, New Mexico
New Mexico
New Mexico is a state located in the southwest and western regions of the United States. New Mexico is also usually considered one of the Mountain States. With a population density of 16 per square mile, New Mexico is the sixth-most sparsely inhabited U.S...
, and Alaska), Canada (Quebec
Quebec
Quebec or is a province in east-central Canada. It is the only Canadian province with a predominantly French-speaking population and the only one whose sole official language is French at the provincial level....
and British Columbia
British Columbia
British Columbia is the westernmost of Canada's provinces and is known for its natural beauty, as reflected in its Latin motto, Splendor sine occasu . Its name was chosen by Queen Victoria in 1858...
), China, Japan, and Korea. The disjunct distribution
Disjunct distribution
In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but widely separated from each other geographically...
of this species in North America and East Asia has been noted to occur in a number of other fungal species as well. Polyozellus multiplex is also found in the Queen Charlotte Islands
Queen Charlotte Islands
Haida Gwaii , formerly the Queen Charlotte Islands, is an archipelago on the North Coast of British Columbia, Canada. Haida Gwaii consists of two main islands: Graham Island in the north, and Moresby Island in the south, along with approximately 150 smaller islands with a total landmass of...
, where it is commercially harvested.
Edibility
Polyozellus multiplex is edibleEdible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...
, and is collected for sale in Asian countries such as Korea, Japan, and China. In North America, it is sometimes collected recreationally, and commercially. The taste is described as mild, and the odor as mild or aromatic. Mycologist David Arora
David Arora
David Arora is an American mycologist, naturalist, and writer. He is the author of two popular books on mushroom identification, Mushrooms Demystified and All That the Rain Promises and More.......
claims the flavor to be inferior to Craterellus. Fruit bodies may be preserved by drying.
Bioactive compounds
Polyozellin
Polyozellin is a chemical which occurs in the mushroom Polyozellus multiplex. It inhibits prolyl endopeptidase , an enzyme that has a role in processing proteins in Alzheimer's disease. Chemicals that inhibit prolyl endopeptidase have attracted research interest due to their potential therapeutic...
—a chemical which can be isolated and purified from P. multiplex—inhibits prolyl endopeptidase
Prolyl endopeptidase
Prolyl endopeptidase or prolyl oligopeptidase, sometimes post-proline cleaving enzyme) is a large cytosolic enzyme that belongs to a distinct class of serine peptidases. It was first described in the cytosol of rabbit brain as an oligopeptidase, which degrades the nonapeptide bradykinin at the...
(PEP), an enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...
that has a role in processing proteins (specifically, amyloid precursor protein
Amyloid precursor protein
Amyloid precursor protein is an integral membrane protein expressed in many tissues and concentrated in the synapses of neurons. Its primary function is not known, though it has been implicated as a regulator of synapse formation, neural plasticity and iron export...
) in Alzheimer's disease
Alzheimer's disease
Alzheimer's disease also known in medical literature as Alzheimer disease is the most common form of dementia. There is no cure for the disease, which worsens as it progresses, and eventually leads to death...
. Chemicals that inhibit PEP have attracted research interest due to their potential therapeutic effects. Further analyses of extracts from P. multiplex revealed similar dibenzofuran
Dibenzofuran
Dibenzofuran, is a heterocyclic organic compound with the chemical structure shown at right. It is an aromatic compound that has two benzene rings fused to one furan ring in the middle. All of the numbered carbon atoms have a hydrogen atom bonded to each of them...
yl derivatives
Derivative (chemistry)
In chemistry, a derivative is a compound that is derived from a similar compound by some chemical or physical process. In the past it was also used to mean a compound that can be imagined to arise from another compound, if one atom is replaced with another atom or group of atoms, but modern...
of polyozellin, each with different chemical properties
Chemical property
A chemical property is any of a material's properties that becomes evident during a chemical reaction; that is, any quality that can be established only by changing a substance's chemical identity...
, including kynapcin
Kynapcin
right|thumb|[[Skeletal formula]] of kynapcin-24Kynapcin is a general name for a number of dibenzofuranyl derivatives of the molecule polyozellin, present in the fungus Polyozellus multiplex. Like polyozellin, it inhibits prolyl endopeptidase, an enzyme that has a role in processing proteins in...
-12, kynapcin-13 and -28, and -24. A total synthesis
Total synthesis
In organic chemistry, a total synthesis is, in principle, the complete chemical synthesis of complex organic molecules from simpler pieces, usually without the aid of biological processes. In practice, these simpler pieces are commercially available in bulk and semi-bulk quantities, and are often...
of kynapcin-24 was achieved in 2009.
Antitumor properties
Research conducted in 2003 suggests that extracts from Polyozellus multiplex may have suppressive effects on stomach cancerStomach cancer
Gastric cancer, commonly referred to as stomach cancer, can develop in any part of the stomach and may spread throughout the stomach and to other organs; particularly the esophagus, lungs, lymph nodes, and the liver...
. The study showed that feeding a low concentration (0.5% or 1%) of the mushroom extract enhanced the activities of the enzymes glutathione S-transferase
Glutathione S-transferase
Enzymes of the glutathione S-transferase family are composed of many cytosolic, mitochondrial, and microsomal proteins. GSTs are present in eukaryotes and in prokaryotes, where they catalyze a variety of reactions and accept endogenous and xenobiotic substrates.GSTs can constitute up to 10% of...
and superoxide dismutase
Superoxide dismutase
Superoxide dismutases are a class of enzymes that catalyze the dismutation of superoxide into oxygen and hydrogen peroxide. As such, they are an important antioxidant defense in nearly all cells exposed to oxygen...
, and increased the abundance of the molecule glutathione
Glutathione
Glutathione is a tripeptide that contains an unusual peptide linkage between the amine group of cysteine and the carboxyl group of the glutamate side-chain...
. The extract also augmented the expression of the protein p53
P53
p53 , is a tumor suppressor protein that in humans is encoded by the TP53 gene. p53 is crucial in multicellular organisms, where it regulates the cell cycle and, thus, functions as a tumor suppressor that is involved in preventing cancer...
. All of these substances protect the human organism against cancer. Additional studies reported in 2004 and 2006 attribute anti-tumor properties to polyozellin.
External links
- California Fungi Photo and further information