Penicillium roqueforti
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''Penicillium roqueforti'' is a common
saprotrophic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ...
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
'' 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 cheeses, flavouring agents, antifungals,
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...
s,
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...
s, and other
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s. The fungus has been a constituent of Roquefort, Stilton, Danish blue, Cabrales, Gorgonzola, and other blue cheeses. Other blue cheeses are made with '' Penicillium glaucum''.


Classification

First described by American mycologist Charles Thom in 1906, ''P. roqueforti'' was initially a heterogeneous species of blue-green, sporulating fungi. They were grouped into different species based on
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological pr ...
differences, but later combined into one species by Kenneth B. Raper and Thom (1949). The ''P. roqueforti'' group got a reclassification in 1996 due to
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bio ...
analysis of
ribosomal DNA Ribosomal DNA (rDNA) is a DNA sequence that codes for ribosomal RNA. These sequences regulate transcription initiation and amplification, and contain both transcribed and non-transcribed spacer segments. In the human genome there are 5 chro ...
sequences. Formerly divided into two varieties―cheese-making (''P. roqueforti'' var. ''roqueforti'') and patulin-making (''P. roqueforti'' var. ''carneum'')―''P. roqueforti'' was reclassified into three species: ''P. roqueforti'', '' P. carneum'', and '' P. paneum''. The complete genome sequence of ''P. roqueforti'' was published in 2014.


Description

As this fungus does not form visible fruiting bodies, descriptions are based on macromorphological characteristics of fungal colonies growing on various standard agar media, and on microscopic characteristics. When grown on Czapek yeast autolysate agar or yeast-extract sucrose (YES) agar, ''P. roqueforti'' colonies are typically 40 mm in diameter, olive brown to dull green (dark green to black on the reverse side of the agar plate), with a velutinous texture. Grown on malt extract agar, colonies are 50 mm in diameter, dull green in color (beige to greyish green on the reverse side), with arachnoid (with many spider-web-like fibers) colony margins. Another characteristic morphological feature of this species is its production of asexual spores in phialides with a distinctive brush-shaped configuration. Evidence for a sexual stage in ''P. roqueforti'' has been found, based in part on the presence of functional mating-type genes and most of the important genes known to be involved in
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
. In 2014, researchers reported inducing the growth of sexual structures in ''P. roqueforti'', including ascogonia,
cleistothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
, and
ascospore An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera ...
s. Genetic analysis and comparison of many different strains isolated from various environments around the world indicate that it is a genetically diverse species. ''P. roqueforti'' is known to be one of the most common spoilage molds of silage. It is also one of several different moulds that can spoil bread.


Uses

The chief industrial use of this species is the production of blue cheeses, such as its namesake Roquefort, Bleu de Bresse, Bleu du Vercors-Sassenage, Brebiblu, Cabrales,
Cambozola Cambozola is a cow's milk cheese that is a combination in style of a French soft-ripened triple cream cheese and Italian Gorgonzola. History Cambozola was patented and industrially produced for the world market by the large German company ...
(Blue Brie),
Cashel Blue ''J&L Grubb'' is an Irish cheese manufacturer, making cows' milk and goats' milk cheeses on their farm near Fethard, South Tipperary. History In 1984, the Grubbs began to make around eight Cashel Blue cheeses at their farm from the milk of their ...
, Danish blue, Polish Rokpol made from cow's milk,
Fourme d'Ambert Fourme d'Ambert is a semi-hard French blue cheese. One of France's oldest cheeses, it dates from as far back as Roman times. It is made from raw cow's milk from the Auvergne region of France, with a distinct, narrow cylindrical shape. The semi- ...
, Fourme de Montbrison, Lanark Blue, Shropshire Blue, and Stilton, and some varieties of Bleu d'Auvergne and Gorgonzola. (Other blue cheeses, including Bleu de Gex and Rochebaron, use '' Penicillium glaucum''.) Strains of the microorganism are also used to produce compounds that can be employed as
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 ...
s, flavours, and fragrances, uses not regulated under the U.S. Toxic Substances Control Act. Its texture is
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous.


Secondary metabolites

Considerable evidence indicates that most strains are capable of producing harmful secondary metabolites (
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s and other mycotoxins) under certain growth conditions. Aristolochene is a sesquiterpenoid compound produced by ''P. roqueforti'', and is likely a precursor to the toxin known as PR toxin, made in large amounts by the fungus. PR-toxin has been implicated in incidents of mycotoxicoses resulting from eating contaminated grains. However, PR toxin is not stable in cheese and breaks down to the less toxic PR imine. Secondary metabolites of ''P. roqueforti'', named andrastins A-D, are found in blue cheese. The andrastins inhibit proteins involved in the efflux of anticancer drugs from multidrug-resistant cancer cells. ''P. roqueforti'' also produces the
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature nerv ...
roquefortine C. However, the levels of roquefortine c in cheese made from it is usually too low to produce toxic effects. The organism can also be used for the production of
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...
s and specialty chemicals, such as
methyl In organic chemistry, a methyl group is an alkyl derived from methane, containing one carbon atom bonded to three hydrogen atoms, having chemical formula . In formulas, the group is often abbreviated as Me. This hydrocarbon group occurs in ...
ketone In organic chemistry, a ketone is a functional group with the structure R–C(=O)–R', where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group –C(=O)– (which contains a carbon-oxygen double b ...
s, including 2-heptanone.


See also

* '' Amaurodon caeruleocaseus'', a fungus named after blue cheese * List of ''Penicillium'' species


References


External links

:''This article is based on text originally fro
a report
of the
United States Environmental Protection Agency The Environmental Protection Agency (EPA) is an independent executive agency of the United States federal government tasked with environmental protection matters. President Richard Nixon proposed the establishment of EPA on July 9, 1970; it ...
.'' {{Taxonbar, from=Q133247 roqueforti Molds used in food production Fungi described in 1906 Taxa named by Charles Thom