Handkea utriformis
Encyclopedia
Handkea utriformis, synonymous
with Lycoperdon utriforme or Calvatia utriformis, is a species of the Lycoperdaceae
family of puffballs. A rather large mushroom
, it may reach dimensions of up to 25 cm (9.8 in) broad by 20 cm (7.9 in) tall. It is commonly known
as the mosaic puffball, a reference to the polygonal-shaped segments the outer surface of the fruiting body develops as it matures. Widespread in northern temperate
zones, it is found frequently on pastures and sandy heath
s, and is edible
when young. H. utriformis has antibiotic
activity against a number of bacteria, and can bioaccumulate
the trace metals copper
and zinc
to relatively high concentrations.
as Lycoperdon utriforme, and since then, has been variously placed in the genera Bovista
, Lycoperdon
, Calvatia
, and Utraria. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia that had distinct microscopic features: Handkea species have a unique type of capillitium (coarse thick-walled hyphae in the gleba
), with curvy slits instead of the usual pores. Although accepted by some authors, the genus concept has been rejected by others.
In the past, the species (when it was known as Calvatia utriformis) has been separated into three varieties (C. utriformis var. utriformis, C. utriformis var. hungarica and C. utriformis var. gruberi) based on differences in the ornamentation of the exoperidium (outer tissue layer of the wall, or peridium
) and spores
. However, a 1997 study of these characters revealed that the three varieties are not clearly demarcated. This study and others suggest that variations in the environmental conditions in which the specimens are grown can affect the development of these characteristics.
Phylogenetic analyses published in 2008 shows that Handkea may be grouped in a clade
along with species from several other genera, including Lycoperdon
, Vascellum, Morganella
, Bovistella
, and Calvatia
. Published in the same year, another DNA analysis of the structure of ITS2
rDNA
transcript
confirmed that H. utriformis is closely related to Lycoperdon echinatum
.
, meaning the spore
s are produced internally, and are only released as the mature fruiting body ages and dries, or is broken.
Young puffballs are typically 6 to 12 cm (2.4 to 4.7 in) across, white, or pale grey-brown; in maturity it may attain dimensions of 25 centimetres (9.8 in) broad by 20 centimetres (7.9 in) tall. The exoperidum is tomentose
—desely covered with a layer of fine matted hairs. The species derives its common name "mosiac puffball" from the mosaic pattern across the top and sides that develops as the fruiting body matures and the outer wall (exoperidium) breaks up into polygonal patches. The underside of the puffball is attached to the ground by a root-like assemblage of hyphae called a rhizomorph
. It is squat in appearance and roughly pear-shaped, not usually taller than it is wide. The flesh (gleba
, or spore bearing mass) is white when young, but becomes brown and powdery upon maturity. The upper skin eventually disintegrates weeks or even months after the puffball's appearance, and the brown spores are released into the air; this process is often hastened by rain, or by being trodden on by cattle. Eventually, all that remains is the sterile cup-shaped base, which can sometimes hold water. This feature may have been the source of the specific epithet, as utrarius is Latin for ‘water carrier’.
s of H. utriformis are roughly spherical, smooth, and thick-walled, with a single oil droplet. They have dimensions of 4.5–5.5 µm
.
s, C. booniana, and C. pachyderma. Calvatia cyathiformis has a purple-colored gleba with a smooth exoperidium; C. booniana has an exoperidium that resembles felt or has tufts of soft "hairs" like H. utriformis but does not have any stem
and has a capillitium with rounded rather than sinuous pits; C. pachyderma has an exoperidium that is thicker and smoother than H. utriformis.
, and is often found in coastal regions. It typically fruits in summer through late autumn.
This fairly large puffball
is edible only when the spore bearing flesh is young, and white. It is said to lack texture, and the taste and odor of the young fruiting bodies are described as "not distinctive". A 2007 study of the fatty acid
composition of various edible Lycoperdaceae species determined the lipid
content of H. utriformis to be quite low, approximately 1.8% (wet weight). The fatty acid content was largely linoleic acid
(42.4%), oleic acid
(23%), palmitic acid
(12.2%), and stearic acid
(3.6%); 17 other fatty acids of various chain lengths and degrees of unsaturation
contributed to the total fatty acid composition.
activity of several Lycoperdaceae revealed that Handkea utriformis has "significantly active" against a number of bacteria
, including Bacillus subtilis
, Escherichia coli
, Klebsiella pneumonia, Pseudomonas aeruginosa
, Salmonella typhimurium, Staphylococcus aureus
, Streptococcus pyogenes
, and Mycobacterium smegmatis
. On the other hand, H. utriformis has low antifungal
activity against the species Candida albicans
, Rhodotorula rubra, and Kluyveromyces fragilis.
and zinc
concentrations in 28 different species of edible mushroom showed that H. utriformis selectively bioaccumulated
both copper (251.9 mg of copper per kilogram of mushroom) and zinc (282.1 mg Zn/kg mushroom) to higher levels than all other mushrooms tested. The authors note that although these trace elements
are important nutritional requirements for humans, and that H. utriformis may be considered a good source of these elements, it is known that absorption of the elements (bioavailability
) from mushrooms is "low due to limited absorption from the small intestine".
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
with Lycoperdon utriforme or Calvatia utriformis, is a species of the Lycoperdaceae
Lycoperdaceae
Lycoperdaceae is a family of approximately 150 fungi now known to lie in the Agaricales. Historically they were placed in their own order Lycoperdales. Members of the Lycoperdaceae family are known as the true puffballs. Unlike other types of fungi that hold spores in gills or teeth, puffballs...
family of puffballs. A rather large 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...
, it may reach dimensions of up to 25 cm (9.8 in) broad by 20 cm (7.9 in) tall. It is commonly known
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...
as the mosaic puffball, a reference to the polygonal-shaped segments the outer surface of the fruiting body develops as it matures. Widespread in northern temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...
zones, it is found frequently on pastures and sandy heath
Heath (habitat)
A heath or heathland is a dwarf-shrub habitat found on mainly low quality acidic soils, characterised by open, low growing woody vegetation, often dominated by plants of the Ericaceae. There are some clear differences between heath and moorland...
s, and is 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...
when young. H. utriformis has antibiotic
Antibiotic
An antibacterial is a compound or substance that kills or slows down the growth of bacteria.The term is often used synonymously with the term antibiotic; today, however, with increased knowledge of the causative agents of various infectious diseases, antibiotic has come to denote a broader range of...
activity against a number of bacteria, and can bioaccumulate
Bioaccumulation
Bioaccumulation refers to the accumulation of substances, such as pesticides, or other organic chemicals in an organism. Bioaccumulation occurs when an organism absorbs a toxic substance at a rate greater than that at which the substance is lost...
the trace metals copper
Copper
Copper is a chemical element with the symbol Cu and atomic number 29. It is a ductile metal with very high thermal and electrical conductivity. Pure copper is soft and malleable; an exposed surface has a reddish-orange tarnish...
and zinc
Zinc
Zinc , or spelter , is a metallic chemical element; it has the symbol Zn and atomic number 30. It is the first element in group 12 of the periodic table. Zinc is, in some respects, chemically similar to magnesium, because its ion is of similar size and its only common oxidation state is +2...
to relatively high concentrations.
Taxonomy
This puffball was originally described in 1790 by French botanist Jean Baptiste François Pierre BulliardJean Baptiste François Pierre Bulliard
Jean Baptiste François Pierre Bulliard was a French physician and botanist....
as Lycoperdon utriforme, and since then, has been variously placed in the genera Bovista
Bovista
Bovista is a genus of fungi in the family Agaricaceae, commonly known as the true puffballs. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the species of Bovista are...
, Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...
, Calvatia
Calvatia
Calvatia is a genus of puffball mushrooms which includes the spectacular giant puffball C. gigantea. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the puffballs,...
, and Utraria. In 1989, German mycologist Hanns Kreisel described the genus Handkea to include species of Calvatia that had distinct microscopic features: Handkea species have a unique type of capillitium (coarse thick-walled hyphae in the gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...
), with curvy slits instead of the usual pores. Although accepted by some authors, the genus concept has been rejected by others.
In the past, the species (when it was known as Calvatia utriformis) has been separated into three varieties (C. utriformis var. utriformis, C. utriformis var. hungarica and C. utriformis var. gruberi) based on differences in the ornamentation of the exoperidium (outer tissue layer of the wall, or peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...
) and spores
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...
. However, a 1997 study of these characters revealed that the three varieties are not clearly demarcated. This study and others suggest that variations in the environmental conditions in which the specimens are grown can affect the development of these characteristics.
Phylogenetic analyses published in 2008 shows that Handkea may be grouped in a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
along with species from several other genera, including Lycoperdon
Lycoperdon
Lycoperdon is a genus of puffball mushrooms. The genus has a widespread distribution and contains about 50 species. In general, it contains the smaller species such as the pear shaped puffball and the gem studded puffball. Most of the time they grow from dead wood and if they grow through the...
, Vascellum, Morganella
Morganella (fungi)
Morganella is a genus of fungi in the family Agaricaceae. The widely distributed genus is prevalent in tropical areas, and contains nine species....
, Bovistella
Bovistella
Bovistella is a genus of fungi in the family Agaricaceae.-External links:*...
, and Calvatia
Calvatia
Calvatia is a genus of puffball mushrooms which includes the spectacular giant puffball C. gigantea. It was formerly classified within the now-obsolete order Lycoperdales, which, following a restructuring of fungal taxonomy brought about by molecular phylogeny, has been split; the puffballs,...
. Published in the same year, another DNA analysis of the structure of ITS2
Internal transcribed spacer
ITS refers to a piece of non-functional RNA situated between structural ribosomal RNAs on a common precursor transcript. Read from 5' to 3', this polycistronic rRNA precursor transcript contains the 5' external transcribed sequence , 18S rRNA, ITS1, 5.8S rRNA, ITS2, 28S rRNA and finally the 3'ETS...
rDNA
Ribosomal DNA
Ribosomal DNA codes for ribosomal RNA. The ribosome is an intracellular macromolecule that produces proteins or polypeptide chains. The ribosome itself consists of a composite of proteins and RNA. As shown in the figure, rDNA consists of a tandem repeat of a unit segment, an operon, composed of...
transcript
Transcription (genetics)
Transcription is the process of creating a complementary RNA copy of a sequence of DNA. Both RNA and DNA are nucleic acids, which use base pairs of nucleotides as a complementary language that can be converted back and forth from DNA to RNA by the action of the correct enzymes...
confirmed that H. utriformis is closely related to Lycoperdon echinatum
Lycoperdon echinatum
Lycoperdon echinatum, commonly known as the spiny puffball or the spring puffball, is a type of puffball mushroom in the genus Lycoperdon. The saprobic species has been found in Africa, Europe, Central America and North America, where it grows on soil in deciduous woods, glades, and pastures...
.
Description
Like all puffballs, Handkea utriformis has a gasteroid basidiocarpBasidiocarp
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...
, meaning the 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...
s are produced internally, and are only released as the mature fruiting body ages and dries, or is broken.
Tomentose
Tomentose is a term used to describe plant hairs that are flattened and matted, forming a woolly coating known as tomentum. Often the hairs are silver or gray-colored...
—desely covered with a layer of fine matted hairs. The species derives its common name "mosiac puffball" from the mosaic pattern across the top and sides that develops as the fruiting body matures and the outer wall (exoperidium) breaks up into polygonal patches. The underside of the puffball is attached to the ground by a root-like assemblage of hyphae called a rhizomorph
Mycelial cord
Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae...
. It is squat in appearance and roughly pear-shaped, not usually taller than it is wide. The flesh (gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...
, or spore bearing mass) is white when young, but becomes brown and powdery upon maturity. The upper skin eventually disintegrates weeks or even months after the puffball's appearance, and the brown spores are released into the air; this process is often hastened by rain, or by being trodden on by cattle. Eventually, all that remains is the sterile cup-shaped base, which can sometimes hold water. This feature may have been the source of the specific epithet, as utrarius is Latin for ‘water carrier’.
Microscopic features
The sporeSpore
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...
s of H. utriformis are roughly spherical, smooth, and thick-walled, with a single oil droplet. They have dimensions of 4.5–5.5 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...
.
Similar species
A number of puffball species resemble H. utriformis, including 'Calvatia cyathiformisCalvatia cyathiformis
Calvatia cyathiformis, or Purple-spored Puffball, is a large edible saprobic species of Calvatia. This terrestrial puffball, has purplish or purple-brown spores, which distinguish it from other large Lycoperdales. It is found mostly in prairie or grasslands of North America, Australia, and probably...
s, C. booniana, and C. pachyderma. Calvatia cyathiformis has a purple-colored gleba with a smooth exoperidium; C. booniana has an exoperidium that resembles felt or has tufts of soft "hairs" like H. utriformis but does not have any 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...
and has a capillitium with rounded rather than sinuous pits; C. pachyderma has an exoperidium that is thicker and smoother than H. utriformis.
Distribution and habitat
Handkea utriformis is widespread, and frequent in northern temperate zones. It is found in Europe, continental Asia, Japan, eastern atlantic North America, Mexico, and South Africa. It has also been collected in Chile, and New Zealand. Growing alone or in small groups, it favors sandy open pastures, or heathsHeath (habitat)
A heath or heathland is a dwarf-shrub habitat found on mainly low quality acidic soils, characterised by open, low growing woody vegetation, often dominated by plants of the Ericaceae. There are some clear differences between heath and moorland...
, and is often found in coastal regions. It typically fruits in summer through late autumn.
Edibility
Puffball
A puffball is a member of any of several groups of fungus in the division Basidiomycota. The puffballs were previously treated as a taxonomic group called the Gasteromycetes or Gasteromycetidae, but they are now known to be a polyphyletic assemblage. The distinguishing feature of all puffballs is...
is edible only when the spore bearing flesh is young, and white. It is said to lack texture, and the taste and odor of the young fruiting bodies are described as "not distinctive". A 2007 study of the fatty acid
Fatty acid
In chemistry, especially biochemistry, a fatty acid is a carboxylic acid with a long unbranched aliphatic tail , which is either saturated or unsaturated. Most naturally occurring fatty acids have a chain of an even number of carbon atoms, from 4 to 28. Fatty acids are usually derived from...
composition of various edible Lycoperdaceae species determined the lipid
Lipid
Lipids constitute a broad group of naturally occurring molecules that include fats, waxes, sterols, fat-soluble vitamins , monoglycerides, diglycerides, triglycerides, phospholipids, and others...
content of H. utriformis to be quite low, approximately 1.8% (wet weight). The fatty acid content was largely linoleic acid
Linoleic acid
Linoleic acid is an unsaturated n-6 fatty acid. It is a colorless liquid at room temperature. In physiological literature, it has a lipid number of 18:2...
(42.4%), oleic acid
Oleic acid
Oleic acid is a monounsaturated omega-9 fatty acid found in various animal and vegetable fats. It has the formula CH37CH=CH7COOH. It is an odorless, colourless oil, although commercial samples may be yellowish. The trans isomer of oleic acid is called elaidic acid...
(23%), palmitic acid
Palmitic acid
Palmitic acid, or hexadecanoic acid in IUPAC nomenclature, is one of the most common saturated fatty acids found in animals and plants. Its molecular formula is CH314CO2H. As its name indicates, it is a major component of the oil from palm trees . Palmitate is a term for the salts and esters of...
(12.2%), and stearic acid
Stearic acid
Stearic acid is the saturated fatty acid with an 18 carbon chain and has the IUPAC name octadecanoic acid. It is a waxy solid, and its chemical formula is CH316CO2H. Its name comes from the Greek word στέαρ "stéatos", which means tallow. The salts and esters of stearic acid are called stearates...
(3.6%); 17 other fatty acids of various chain lengths and degrees of unsaturation
Unsaturated compound
In organic chemistry, a saturated compound is a chemical compound that has of a chain of carbon atoms linked together by single bonds and has hydrogen atoms filling all of the other bonding orbitals of the carbon atoms. Alkanes are an example of saturated compounds...
contributed to the total fatty acid composition.
Antibiotic activity
A 2005 study of the antimicrobialAntimicrobial
An anti-microbial is a substance that kills or inhibits the growth of microorganisms such as bacteria, fungi, or protozoans. Antimicrobial drugs either kill microbes or prevent the growth of microbes...
activity of several Lycoperdaceae revealed that Handkea utriformis has "significantly active" against a number of bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, including Bacillus subtilis
Bacillus subtilis
Bacillus subtilis, known also as the hay bacillus or grass bacillus, is a Gram-positive, catalase-positive bacterium commonly found in soil. A member of the genus Bacillus, B. subtilis is rod-shaped, and has the ability to form a tough, protective endospore, allowing the organism to tolerate...
, Escherichia coli
Escherichia coli
Escherichia coli is a Gram-negative, rod-shaped bacterium that is commonly found in the lower intestine of warm-blooded organisms . Most E. coli strains are harmless, but some serotypes can cause serious food poisoning in humans, and are occasionally responsible for product recalls...
, Klebsiella pneumonia, Pseudomonas aeruginosa
Pseudomonas aeruginosa
Pseudomonas aeruginosa is a common bacterium that can cause disease in animals, including humans. It is found in soil, water, skin flora, and most man-made environments throughout the world. It thrives not only in normal atmospheres, but also in hypoxic atmospheres, and has, thus, colonized many...
, Salmonella typhimurium, Staphylococcus aureus
Staphylococcus aureus
Staphylococcus aureus is a facultative anaerobic Gram-positive coccal bacterium. It is frequently found as part of the normal skin flora on the skin and nasal passages. It is estimated that 20% of the human population are long-term carriers of S. aureus. S. aureus is the most common species of...
, Streptococcus pyogenes
Streptococcus pyogenes
Streptococcus pyogenes is a spherical, Gram-positive bacterium that is the cause of group A streptococcal infections. S. pyogenes displays streptococcal group A antigen on its cell wall. S...
, and Mycobacterium smegmatis
Mycobacterium smegmatis
Mycobacterium smegmatis is 3.0 to 5.0 µm long with a bacillus shape, an acid-fast bacterial species in the phylum Actinobacteria. It can be stained by Ziehl-Neelsen method and the auramine-rhodamine fluorescent method. It was first reported in November 1884 by Lustgarten, who found a bacillus...
. On the other hand, H. utriformis has low antifungal
Antifungal
* Antifungal medication, a medication used to treat fungal infection s such as athlete's foot , ringworm, candidiasis, etc.* Antifungal protein, a protein family* an adjective referring to a fungicide compound...
activity against the species Candida albicans
Candida albicans
Candida albicans is a diploid fungus that grows both as yeast and filamentous cells and a causal agent of opportunistic oral and genital infections in humans. Systemic fungal infections including those by C...
, Rhodotorula rubra, and Kluyveromyces fragilis.
Bioaccumulation
A study of the copperCopper
Copper is a chemical element with the symbol Cu and atomic number 29. It is a ductile metal with very high thermal and electrical conductivity. Pure copper is soft and malleable; an exposed surface has a reddish-orange tarnish...
and zinc
Zinc
Zinc , or spelter , is a metallic chemical element; it has the symbol Zn and atomic number 30. It is the first element in group 12 of the periodic table. Zinc is, in some respects, chemically similar to magnesium, because its ion is of similar size and its only common oxidation state is +2...
concentrations in 28 different species of edible mushroom showed that H. utriformis selectively bioaccumulated
Bioaccumulation
Bioaccumulation refers to the accumulation of substances, such as pesticides, or other organic chemicals in an organism. Bioaccumulation occurs when an organism absorbs a toxic substance at a rate greater than that at which the substance is lost...
both copper (251.9 mg of copper per kilogram of mushroom) and zinc (282.1 mg Zn/kg mushroom) to higher levels than all other mushrooms tested. The authors note that although these trace elements
Micronutrient
Micronutrients are nutrients required by humans and other living things throughout life in small quantities to orchestrate a whole range of physiological functions, but which the organism itself cannot produce. For people, they include dietary trace minerals in amounts generally less than 100...
are important nutritional requirements for humans, and that H. utriformis may be considered a good source of these elements, it is known that absorption of the elements (bioavailability
Bioavailability
In pharmacology, bioavailability is a subcategory of absorption and is used to describe the fraction of an administered dose of unchanged drug that reaches the systemic circulation, one of the principal pharmacokinetic properties of drugs. By definition, when a medication is administered...
) from mushrooms is "low due to limited absorption from the small intestine".
External links
- Mycobank
- Medicinal Mushrooms Medicinal properties and bioactive compounds