Penicillium Vermiculatum
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''Penicillium'' () is a genus of
ascomycetous Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The de ...
fungi that is part of the
mycobiome The mycobiome, mycobiota, or fungal microbiome, is the fungi, fungal community in and on an organism. The word “mycobiome” comes from the ancient Greek μύκης (mukēs), meaning "fungus" with the suffix “biome” derived from the Greek ...
of many species and is of major importance in the natural environment, in food spoilage, and in food and drug production. Some members of the genus produce
penicillin Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
, a molecule that is used as an
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
, which kills or stops the growth of certain kinds of bacteria. Other species are used in
cheesemaking Cheesemaking (or caseiculture) is the craft of making cheese. The production of cheese, like many other food preservation processes, allows the nutritional and economic value of a food material, in this case milk, to be preserved in concentrat ...
. According to the ''Dictionary of the Fungi'' (10th edition, 2008), the widespread genus contains over 300 species.


Taxonomy

The genus was first described in the scientific literature by
Johann Heinrich Friedrich Link Johann Heinrich Friedrich Link (2 February 1767 – 1 January 1851) was a German naturalist and botanist. Biography Link was born at Hildesheim as a son of the minister August Heinrich Link (1738–1783), who taught him love of nature throug ...
in his 1809 work ''Observationes in ordines plantarum naturales''; he wrote, "''Penicillium. Thallus e floccis caespitosis septatis simplicibus aut ramosis fertilibus erectis apice penicillatis''", where ''penicillatis'' means "having tufts of fine hair". Link included three species—'' P. candidum'', '' P. expansum'', and '' P. glaucum''—all of which produced a brush-like conidiophore (asexual spore-producing structure). The common apple rot fungus ''P. expansum'' was later selected as the type species. In his 1979
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
, John I. Pitt divided ''Penicillium'' into four
subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
based on conidiophore morphology and branching pattern: ''Aspergilloides'', ''Biverticillium'', ''Furcatum'', and ''Penicillium''. Species included in subgenus ''Biverticillium'' were later merged into '' Talaromyces''.


Species

Selected species include; * '' Penicillium albocoremium'' * '' Penicillium aurantiogriseum'', a grain contaminant * '' Penicillium bilaiae'', which is an agricultural inoculant * ''
Penicillium camemberti ''Penicillium camemberti'' is a species of fungus in the genus ''Penicillium''. It is used in the production of Camembert, Brie, Langres, Coulommiers, and Cambozola cheeses, on which colonies of ''P. camemberti'' form a hard, white crust. I ...
'', which is used in the production of Camembert and Brie
cheese Cheese is a dairy product produced in wide ranges of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk, usually the milk of cows, buffalo, goats, or sheep. During production, ...
s * ''
Penicillium candidum ''Penicillium camemberti'' is a species of fungus in the genus ''Penicillium''. It is used in the production of Camembert, Brie, Langres, Coulommiers, and Cambozola cheeses, on which colonies of ''P. camemberti'' form a hard, white crust. ...
'', which is used in making Brie and Camembert. It has been reduced to synonymy with ''Penicillium camemberti'' * '' Penicillium chrysogenum'' (previously known as '' Penicillium notatum''), which produces the
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
penicillin Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
* ''
Penicillium claviforme ''Penicillium claviforme'' is a species of ''Penicillium'' within the phylum Ascomycota. *It is found within the subgenus ''Terverticillium''. *It has a coremium-type morphology in the sexual structures, named for its resemblance to matchsticks. ...
'' * ''
Penicillium commune ''Penicillium commune'' is an indoor fungus belonging to the genus ''Penicillium''. It is known as one of the most common fungi spoilage moulds on cheese. It also grows on and spoils other foods such as meat products and fat-containing products ...
'' * ''
Penicillium crustosum ''Penicillium crustosum'' is a blue-green or blue-grey mold that can cause food spoilage, particularly of protein-rich foods such as meats and cheeses. It is identified by its complex biseriate conidiophores on which phialides produce asexual spo ...
'' * '' Penicillium digitatum'', a ''Citrus'' pathogen * ''
Penicillium echinulatum ''Penicillium echinulatum'' is a mold species in the genus ''Penicillium''. It is a source of cellulase. 5-Hydroxymaltol and mycophenolic acid Mycophenolic acid (MPA) is an immunosuppressant medication used to prevent rejection following o ...
'' produces Mycophenolic acid * '' Penicillium expansum'', a pathogen of apples and other fruit, produces patulin * ''
Penicillium glabrum ''Penicillium glabrum'' is a plant pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be refer ...
'' * '' Penicillium imranianum'' * ''
Penicillium italicum ''Penicillium italicum'' is a plant pathogen. It is a common post harvest disease commonly associated with citrus fruits. Management Inoculation of healthy fruit can be diminished and controlled by careful picking, handling, and packaging of the ...
'', a ''Citrus'' pathogen * '' Penicillium lacussarmientei'' * '' Penicillium lusitanum'', isolated from marine habitat * ''
Penicillium purpurogenum ''Penicillium purpurogenum'' is a plant pathogen infecting strawberries. It produces rubratoxin B, a mycotoxin with anticarcinogenic properties, as well as monascus pigments. See also * List of strawberry diseases This article is a list of d ...
'' * ''
Penicillium roqueforti ''Penicillium roqueforti'' is a common saprotrophic fungus in the genus ''Penicillium''. Widespread in nature, it can be isolated from soil, decaying organic matter, and plants. The major industrial use of this fungus is the production of blue ...
'', which is used in making Roquefort,
Danish Blue cheese Danablu, often marketed under the trademark Danish Blue Cheese within North America, is a strong, blue-veined cheese. This semi-soft creamery cheese is typically drum- or block-shaped and has a yellowish, slightly moist, edible rind. Made from f ...
, English Blue
Stilton cheese Stilton is an English cheese, produced in two varieties: Blue, which has '' Penicillium roqueforti'' added to generate a characteristic smell and taste, and White, which does not. Both have been granted the status of a protected designation of o ...
, and
Gorgonzola cheese Gorgonzola (; ) is a veined blue cheese, originally from Italy, made from unskimmed cow's milk. It can be buttery or firm, crumbly and quite salty, with a "bite" from its blue veining. History Historically, gorgonzola has been produced for ...
* ''
Penicillium stoloniferum ''Penicillium brevicompactum'' is a mould species in the genus ''Penicillium''. Mycophenolic acid Mycophenolic acid (MPA) is an immunosuppressant medication used to prevent rejection following organ transplantation and to treat autoimmune c ...
'' * '' Penicillium ulaiense'', a ''Citrus'' pathogen in Asia * '' Penicillium verrucosum'', a grain contaminant which produces ochratoxin A * '' Penicillium viridicatum''


Etymology

The genus name is derived from the Latin root ''penicillum'', meaning "painter's brush", and refers to the chains of conidia that resemble a broom.


Characteristics

The thallus ( mycelium) consists of highly branched networks of multinucleated, usually colourless
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
, with each pair of cells separated by a
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatri ...
. Conidiophores are at the end of each branch accompanied by green spherical constricted units called
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
. These propagules play a significant role in reproduction; conidia are the main dispersal strategy of these fungi. Sexual reproduction involves the production of ascospores, commencing with the fusion of an
archegonium An archegonium (pl: archegonia), from the ancient Greek ''ἀρχή'' ("beginning") and ''γόνος'' ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female ga ...
and an antheridium, with sharing of nuclei. The irregularly distributed
asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...
contain eight unicellular ascospores each.


Ecology

Species of ''Penicillium'' are ubiquitous soil fungi preferring cool and moderate climates, commonly present wherever organic material is available. Saprophytic species of ''Penicillium'' and '' Aspergillus'' are among the best-known representatives of the
Eurotiales The Eurotiales are an order of sac fungi, also known as the green and blue molds. It was circumscribed in 1980. Classification Currently the order Eurotiales contains 5 families, 28 genera and 1280 species: *Family Aspergillaceae Monascaceae ...
and live mainly on organic biodegradable substances. Commonly known in America as molds, they are among the main causes of food spoilage, especially species of
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
''Penicillium''. Many species produce highly toxic mycotoxins. The ability of these ''Penicillium'' species to grow on seeds and other stored foods depends on their propensity to thrive in low humidity and to colonize rapidly by aerial dispersion while the seeds are sufficiently moist. Some species have a blue color, commonly growing on old bread and giving it a blue fuzzy texture. Some ''Penicillium'' species affect the fruits and bulbs of plants, including '' P. expansum'', apples and pears; '' P. digitatum'', citrus fruits; and '' P. allii'', garlic. Some species are known to be pathogenic to animals; '' P. corylophilum'', '' P. fellutanum'', '' P. implicatum'', '' P. janthinellum'', '' P. viridicatum'', and '' P. waksmanii'' are potential pathogens of mosquitoes. ''Penicillium'' species are present in the air and dust of indoor environments, such as homes and public buildings. The fungus can be readily transported from the outdoors, and grow indoors using building material or accumulated soil to obtain nutrients for growth. ''Penicillium'' growth can still occur indoors even if the relative humidity is low, as long as there is sufficient moisture available on a given surface. A British study determined that ''Aspergillus''- and ''Penicillium''-type spores were the most prevalent in the indoor air of residential properties, and exceeded outdoor levels. Even ceiling tiles can support the growth of ''Penicillium''—as one study demonstrated—if the relative humidity is 85% and the moisture content of the tiles is greater than 2.2%. Some ''Penicillium'' species cause damage to machinery and the combustible materials and lubricants used to run and maintain them. For example, '' P. chrysogenum (formerly P. notatum)'', '' P. steckii'', ''   P. cyclopium'', and '' P. nalgiovensis'' affect fuels; ''P. chrysogenum'', '' P. rubrum'', and '' P. verrucosum'' cause damage to oils and lubricants; '' P. regulosum'' damages optical and protective glass.


Economic value

Several species of the genus ''Penicillium'' play a central role in the production of cheese and of various meat products. To be specific, ''Penicillium'' molds are found in Blue cheese. ''
Penicillium camemberti ''Penicillium camemberti'' is a species of fungus in the genus ''Penicillium''. It is used in the production of Camembert, Brie, Langres, Coulommiers, and Cambozola cheeses, on which colonies of ''P. camemberti'' form a hard, white crust. I ...
'' and ''
Penicillium roqueforti ''Penicillium roqueforti'' is a common saprotrophic fungus in the genus ''Penicillium''. Widespread in nature, it can be isolated from soil, decaying organic matter, and plants. The major industrial use of this fungus is the production of blue ...
'' are the molds on Camembert, Brie, Roquefort, and many other cheeses. ''
Penicillium nalgiovense ''Penicillium nalgiovense'' is an anamorph species of the genus ''Penicillium'' with lipolytic and proteolytic Proteolysis is the breakdown of proteins into smaller polypeptides or amino acids. Uncatalysed, the hydrolysis of peptide bonds is e ...
'' is used in soft mold-ripened cheeses, such as Nalžovy (ellischau) cheese, and to improve the taste of sausages and hams, and to prevent colonization by other molds and bacteria. In addition to their importance in the food industry, species of ''Penicillium'' and ''Aspergillus'' serve in the production of a number of biotechnologically produced enzymes and other macromolecules, such as gluconic, citric, and tartaric acids, as well as several
pectinase Pectinases are a group of enzymes that breaks down pectin, a polysaccharide found in plant cell walls, through hydrolysis, transelimination and deesterification reactions. Commonly referred to as pectic enzymes, they include pectolyase, pectozym ...
s,
lipase Lipase ( ) is a family of enzymes that catalyzes the hydrolysis of fats. Some lipases display broad substrate scope including esters of cholesterol, phospholipids, and of lipid-soluble vitamins and sphingomyelinases; however, these are usually tr ...
, amylases, cellulases, and proteases. Some ''Penicillium'' species have shown potential for use in bioremediation, more specifically mycoremediation, because of their ability to break down a variety of xenobiotic compounds. The genus includes a wide variety of species molds that are the source molds of major
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention of ...
s.
Penicillin Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
, a drug produced by '' P. chrysogenum (formerly P. notatum),'' was accidentally discovered by
Alexander Fleming Sir Alexander Fleming (6 August 1881 – 11 March 1955) was a Scottish physician and microbiologist, best known for discovering the world's first broadly effective antibiotic substance, which he named penicillin. His discovery in 1928 of w ...
in 1929, and found to inhibit the growth of Gram-positive bacteria (see
beta-lactams A beta-lactam (β-lactam) ring is a four-membered lactam. A ''lactam'' is a cyclic amide, and ''beta''-lactams are named so because the nitrogen atom is attached to the β-carbon atom relative to the carbonyl. The simplest β-lactam possible is ...
). Its potential as an antibiotic was realized in the late 1930s, and
Howard Florey Howard Walter Florey, Baron Florey (24 September 189821 February 1968) was an Australian pharmacologist and pathologist who shared the Nobel Prize in Physiology or Medicine in 1945 with Sir Ernst Chain and Sir Alexander Fleming for his role in ...
and
Ernst Chain Sir Ernst Boris Chain (19 June 1906 – 12 August 1979) was a German-born British biochemist best known for being a co-recipient of the Nobel Prize in Physiology or Medicine for his work on penicillin. Life and career Chain was born in Be ...
purified and concentrated the compound. The drug's success in saving soldiers in World War II who had been dying from infected wounds resulted in Fleming, Florey and Chain jointly winning the Nobel Prize in Medicine in 1945. Griseofulvin is an antifungal drug and a potential chemotherapeutic agent that was discovered in '' P. griseofulvum''. Additional species that produce compounds capable of inhibiting the growth of tumor cells '' in vitro'' include: '' P. pinophilum'', '' P. canescens'', and '' P. glabrum''.


Reproduction

Although many
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s are able to reproduce sexually, as much as 20% of fungal species had been thought to reproduce exclusively by asexual means. However recent studies have revealed that
sex Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones (ova, oft ...
occurs even in some of the supposedly asexual species. For example, sexual capability was recently shown for the fungus ''
Penicillium roqueforti ''Penicillium roqueforti'' is a common saprotrophic fungus in the genus ''Penicillium''. Widespread in nature, it can be isolated from soil, decaying organic matter, and plants. The major industrial use of this fungus is the production of blue ...
'', used as a starter for blue cheese production. This finding was based, in part, on evidence for functional mating type (MAT) genes that are involved in fungal sexual compatibility, and the presence in the sequenced genome of most of the important genes known to be involved in meiosis. '' Penicillium chrysogenum'' is of major medical and historical importance as the original and present-day industrial source of the antibiotic penicillin. The species was considered asexual for more than 100 years despite concerted efforts to induce sexual reproduction. However, in 2013, Bohm et al. finally demonstrated sexual reproduction in ''P. chrysogenum''. These findings with ''Penicillium'' species are consistent with accumulating evidence from studies of other eukaryotic species that sex was likely present in the common ancestor of all
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. Furthermore, these recent results suggest that sex can be maintained even when very little genetic variability is produced. Prior to 2013, when the "
one fungus, one name In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the Biological life cycle, life cycles of fungi in the Phylum, phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a Asc ...
" nomenclature change came into effect, ''Penicillium'' was used as the genus for anamorph (clonal forms) of fungi and ''Talaromyces'' was used for the teleomorph (sexual forms) of fungi. After 2013 however, fungi were reclassified based on their genetic relatedness to each other and now the genera ''Penicillium'' and ''Talaromyces'' both contain some species capable of only clonal reproduction and others that can reproduce sexually.


References

*


External links

* * {{Authority control Medicinal fungi Taxa named by Johann Heinrich Friedrich Link Taxa described in 1809 Eurotiomycetes genera