Lycoperdon perlatum
Encyclopedia
Lycoperdon perlatum, commonly known as the common puffball, warted puffball, gem-studded puffball or devil's snuff-box, is a species of puffball
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...

 mushroom in the Agaricaceae
Agaricaceae
The Agaricaceae are a family of basidiomycete fungi and includes the genus Agaricus, as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. The genus contains 85 genera and 1340 species.-Genera:...

 family. A common species with a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...

, it is a moderate-sized puffball with a round fruit body tapering to a wide stalk, and dimensions of 1.5 to 6 cm (0.590551181102362 to 2.4 in) wide by 3 to 7 cm (1.2 to 2.8 in) tall. It is off-white with a top covered in short spiny bumps or 'jewels', which are easily rubbed off. When mature they become brown and a hole in the top opens to release spores in a burst when the body is compressed by raindrops, a touch, falling nuts, etc. It grows in fields, gardens, and along roadsides, as well as in grassy clearings in woods. The puffball 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 and the internal flesh is completely white. Laboratory tests indicate that extracts of the puffball have antimicrobial
Antimicrobial
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...

 and antifungal activities.

Taxonomy

The species was first described in the scientific literature in 1796 by mycologist Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

. The variant Lycoperdon gemmatum var. perlatum, published by Elias Magnus Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

 in 1829, is a synonym. Some authorities consider Lycoperdon gemmatum, described by August Batsch
August Batsch
Dr August Johann Georg Karl Batsch was a German naturalist. He was a recognised authority on mushrooms, and also described new species of ferns, bryophytes, and seed plants.- Life and career :...

 in 1783, to be a synonym.

The specific epithet perlatum is Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 for "widespread". It is 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 common puffball, the gem-studded puffball (or gemmed puffball), the warted puffball, or the devil's snuff-box.

Description

The fruit body ranges in shape from pear-like with a flattened top, to nearly spherical, and reaches dimensions of 1.5 to 6 cm (0.590551181102362 to 2.4 in) wide by 3 to 7 cm (1.2 to 2.8 in) tall. It has a stem-like base. The outer surface of the fruit body (the exoperidium
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:...

) is covered in short cone-shaped spines that are interspersed with granular warts. The spines, which are whitish, gray, or brown, can be easily rubbed off, and leave reticulate pock marks or scars after they are removed. The base of the puffball is thick, and has internal chambers. It is initially white, but turns yellow, olive, or brownish in age. The reticulate pattern resulting from the rubbed-off spines is less evident on the base. In maturity, the exoperidium at the top of the puffball sloughs away, revealing a pre-formed hole (ostiole
Ostiole
An ostiole is a small hole or opening through which algae or ascomycetal fungi release their mature spores. The term is also used in higher plants, for example to denote the opening of the involuted fig inflorescence through which fig wasps enter to pollinate and breed....

) in the endoperidium, through which the spores can escape. In young puffballs, the internal contents, 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...

, is white and firm, but turns brown and powdery as 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 mature. The gleba contains minute chambers that are lined with 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...

 (the fertile, 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...

-bearing tissue); the chambers collapse when the spores mature.

The spores are spherical, thick-walled, covered with minute spines, and measure 3.5–4.5 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...

 in diameter. The capillitia (threadlike filaments in the gleba in which spores are embedded) are yellow-brown to brownish in color, lack septae
Septum
In anatomy, a septum is a wall, dividing a cavity or structure into smaller ones.-In human anatomy:...

, and measure 3–7.5 μm in diameter. 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...

 (spore-bearing cells) are club-shaped, four-spored, and measure 7–9 by 4–5 μm. The basidia bear four slender sterigmata of unequal length ranging from 5–10 μm long. The surface spines are made of chains of pseudoparenchymatous hyphae (resembling the parenchyma
Parenchyma
Parenchyma is a term used to describe a bulk of a substance. It is used in different ways in animals and in plants.The term is New Latin, f. Greek παρέγχυμα - parenkhuma, "visceral flesh", f. παρεγχεῖν - parenkhein, "to pour in" f. para-, "beside" + en-, "in" + khein, "to pour"...

  of higher plants), in which the individual hyphal cells are spherical to elliptical in shape, thick-walled (up to 1 μm), and measure 13–40 by 9–35 μm. These hyphae do not have 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.

Edibility

Lycoperdon perlatum is considered to be a good edible mushroom
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, when 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...

 is still homogeneous and white. They have been referred to as "poor man's sweetbread" due to their texture and flavor. The fruit bodies can be eaten by slicing and frying in batter or egg and breadcrumbs, or used in soups. The puffballs become inedible as they mature: the gleba becomes yellow-tinged then finally develops into a mass of powdery olive-green spores.

The immature 'buttons' or 'eggs' of deadly Amanita
Amanita
The genus Amanita contains about 600 species of agarics including some of the most toxic known mushrooms found worldwide. This genus is responsible for approximately 95% of the fatalities resulting from mushroom poisoning, with the death cap accounting for about 50% on its own...

species can be confused with puffballs. For this reason puffballs being collected for the table should always be sliced vertically and inspected for the internal developing structures of a mushroom. Amanita
Amanita
The genus Amanita contains about 600 species of agarics including some of the most toxic known mushrooms found worldwide. This genus is responsible for approximately 95% of the fatalities resulting from mushroom poisoning, with the death cap accounting for about 50% on its own...

s will generally not have 'jewels' or a bumpy external surface.

The spores are ornamented with many sharp microscopic spines and can cause severe irritation of the lung (lycoperdonosis
Lycoperdonosis
Lycoperdonosis is a respiratory disease caused by the inhalation of large quantities of spores from mature puffballs. It is classified as a hypersensitivity pneumonitis —an inflammation of the alveoli within the lung caused by hypersensitivity to inhaled organic dusts...

) when deliberately inhaled. This condition has been reported to afflict dogs that play or run where puffballs are present.

Similar species

There are several other puffball species with which L. perlatum might be confused. L. nettyanum, found in the Pacific Northwest
Pacific Northwest
The Pacific Northwest is a region in northwestern North America, bounded by the Pacific Ocean to the west and, loosely, by the Rocky Mountains on the east. Definitions of the region vary and there is no commonly agreed upon boundary, even among Pacific Northwesterners. A common concept of the...

 region of the United States, is covered in granular patches, but these granules adhere more strongly to the surface than those of L. perlatum. L. pyriforme lacks prominent spines on the surface, and grows on rotting wood—although if growing on buried wood, it may appear to be terrestrial. The widely distributed and common L. umbrinum
Lycoperdon umbrinum
Lycoperdon umbrinum, commonly known as the umber-brown puffball, is a type of Puffball mushroom in the genus Lycoperdon. It is found in China,, Europe, and North America.-Description:...

has spines that do not leave scars when rubbed off, a gleba that varies in color from dark brown to purple-brown at maturity, and a purple-tinged base. The small and rare species L. muscorum grows in deep moss. L. peckii can be distinguished from L. pyriforme by the lavender-tinged spines it has when young. L. rimulatum has purplish spores, and an almost completely smooth exoperidium.

Habitat and distribution

Lycoperdon perlatum grows solitarily, scattered, or in groups or clusters on the ground. It can also grow in fairy ring
Fairy ring
A fairy ring, also known as fairy circle, elf circle, elf ring or pixie ring, is a naturally occurring ring or arc of mushrooms. The rings may grow to over in diameter, and they become stable over time as the fungus grows and seeks food underground. They are found mainly in forested areas, but...

s. Typical habitats include woods, grassy areas, and along roads. It has been reported from Pinus patula
Pinus patula
Patula pine, pino patula, pinus patula is a tree native to the highlands of Mexico. It grows from 24° to 18° North latitude and 1800 to 2700 m above sea level. 30 m tall...

plantations in Tamil Nadu
Tamil Nadu
Tamil Nadu is one of the 28 states of India. Its capital and largest city is Chennai. Tamil Nadu lies in the southernmost part of the Indian Peninsula and is bordered by the union territory of Pondicherry, and the states of Kerala, Karnataka, and Andhra Pradesh...

, India. The puffball sometimes confuses golfers because of its resemblance to a golfball when viewed from a distance. It is widely distributed, and has been reported from Africa (Tanzania
Tanzania
The United Republic of Tanzania is a country in East Africa bordered by Kenya and Uganda to the north, Rwanda, Burundi, and the Democratic Republic of the Congo to the west, and Zambia, Malawi, and Mozambique to the south. The country's eastern borders lie on the Indian Ocean.Tanzania is a state...

), Asia (Himalayas, Japan), Australia, and South America (Brazil). In North America, where it is considered the most common puffball species, it ranges from 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...

 to Mexico.

The puffball bioaccumulates
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...

 heavy metals present in the soil, and can be used as a bioindicator
Bioindicator
Biological indicators are species used to monitor the health of an environment or ecosystem. They are any biological species or group of species whose function, population, or status can be used to determine ecosystem or environmental integrity. An example of such a group are the copepods and other...

 of soil pollution by heavy metals and selenium
Selenium
Selenium is a chemical element with atomic number 34, chemical symbol Se, and an atomic mass of 78.96. It is a nonmetal, whose properties are intermediate between those of adjacent chalcogen elements sulfur and tellurium...

. In one 1977 study, samples collected from grassy areas near the side of an interstate highway in Connecticut
Connecticut
Connecticut is a state in the New England region of the northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, and the state of New York to the west and the south .Connecticut is named for the Connecticut River, the major U.S. river that approximately...

 were shown to have high concentrations of cadmium
Cadmium
Cadmium is a chemical element with the symbol Cd and atomic number 48. This soft, bluish-white metal is chemically similar to the two other stable metals in group 12, zinc and mercury. Similar to zinc, it prefers oxidation state +2 in most of its compounds and similar to mercury it shows a low...

 and lead
Lead
Lead is a main-group element in the carbon group with the symbol Pb and atomic number 82. Lead is a soft, malleable poor metal. It is also counted as one of the heavy metals. Metallic lead has a bluish-white color after being freshly cut, but it soon tarnishes to a dull grayish color when exposed...

.

Chemistry

Several steroid derivatives have been isolated and identified from fruit bodies of L. perlatum, including (S)-23-hydroxylanostrol, ergosterol α-endoperoxide, ergosterol 9,11-dehydroendoperoxide and (23E)-lanosta-8,23-dien-3β,25-diol. The compounds 3-octanone, 1-octen-3-ol
1-Octen-3-ol
1-Octen-3-ol, octenol for short , is a chemical that attracts biting insects such as mosquitos. It is contained in human breath and sweat, and it was once believed that insect repellent DEET works by blocking the insects' octenol odorant receptors.Octenol is used in combination with carbon dioxide...

, and (Z)-3-octen-1-ol are the predominant components of the volatile
Volatility (chemistry)
In chemistry and physics, volatility is the tendency of a substance to vaporize. Volatility is directly related to a substance's vapor pressure. At a given temperature, a substance with higher vapor pressure vaporizes more readily than a substance with a lower vapor pressure.The term is primarily...

 chemicals that give the puffball its odor and flavor. Extracts of the puffball contain relatively high levels of antimicrobial
Antimicrobial
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 against the human pathogenic bacteria 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...

, 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...

, 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...

, and 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...

, with an efficiency comparable to that of ampicillin
Ampicillin
Ampicillin is a beta-lactam antibiotic that has been used extensively to treat bacterial infections since 1961. Until the introduction of ampicillin by the British company Beecham, penicillin therapies had only been effective against Gram-positive organisms such as staphylococci and streptococci...

. These results corroborate an earlier study that additionally reported antibacterial activity against Salmonella typhimurium, 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...

. Extracts of the puffball have also been reported to have antifungal activity against 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...

, Candida tropicalis
Candida tropicalis
Candida tropicalis is a species of yeast in the genus Candida. It is easily recognized as a common medical yeast pathogen, existing as part of the normal human flora.-External links:*...

, Aspergillus fumigatus
Aspergillus fumigatus
Aspergillus fumigatus is a fungus of the genus Aspergillus, and is one of the most common Aspergillus species to cause disease in individuals with an immunodeficiency....

, Alternaria solani
Alternaria solani
Alternaria solani is a fungal pathogen, producing a disease in tomato and potato plants called early blight. The pathogen produces distinctive "bullseye" patterned leaf spots and can also cause stem lesions and fruit rot on tomato and tuber blight on potato. Despite the name "early," foliar...

, Botrytis cinerea
Botrytis cinerea
Botrytis cinerea is a necrotrophic fungus that affects many plant species, although its most notable hosts may be wine grapes. In viticulture, it is commonly known as botrytis bunch rot; in horticulture, it is usually called grey mould or gray mold.The fungus gives rise to two different kinds of...

, and Verticillium dahliae
Verticillium dahliae
Verticillium dahliae is a fungal plant pathogen. It causes verticillium wilt in many plant species, causing leaves to curl and discolor. It may cause death in some plants.- External links :* *...

. A 2009 study found L. perlatum puffballs to contain the phenolic
Natural phenol
Natural phenols, bioavailable phenols, plant phenolics, low molecular weight phenols or phenoloids are a class of natural products. They are small molecules containing one or more phenolic group. These molecules are smaller in size than polyphenols, containing less than 12 phenolic groups...

 compound cinnamic acid
Cinnamic acid
Cinnamic acid is a white crystalline organic acid, which is slightly soluble in water.It is obtained from oil of cinnamon, or from balsams such as storax. It is also found in shea butter and is the best indication of its environmental history and post-extraction conditions...

 at a concentration of about 14 milligrams per kilogram of mushroom. The fruit bodies contain the pigment melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...

.

The unusual amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

 lycoperdic acid 3-(5(S)-carboxy-2-oxotetrahydrofuran-5(S)-yl)-2(S)-alanine was isolated from the mushroom, and reported in a 1979 publication. Based on the structural similarity of the new amino acid with (S)-glutamic acid
Glutamic acid
Glutamic acid is one of the 20 proteinogenic amino acids, and its codons are GAA and GAG. It is a non-essential amino acid. The carboxylate anions and salts of glutamic acid are known as glutamates...

, (S)-(+)-lycoperdic acid is expected to have antagonistic
Receptor antagonist
A receptor antagonist is a type of receptor ligand or drug that does not provoke a biological response itself upon binding to a receptor, but blocks or dampens agonist-mediated responses...

 or agonistic activity for the glutamate receptor
Glutamate receptor
Glutamate receptors are synaptic receptors located primarily on the membranes of neuronal cells. Glutamate is one of the 20 amino acids used to assemble proteins and as a result is abundant in many areas of the body, but it also functions as a neurotransmitter and is particularly abundant in the...

 in the mammalian central nervous system. Methods to synthesize
Organic synthesis
Organic synthesis is a special branch of chemical synthesis and is concerned with the construction of organic compounds via organic reactions. Organic molecules can often contain a higher level of complexity compared to purely inorganic compounds, so the synthesis of organic compounds has...

 the compounds were reported in 1992, 1995, and 2002.

The predominant 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...

s in the puffball are 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...

 (37% of the total fatty acids), 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...

 (24%), 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...

 (14.5%), 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...

(6.4%).
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