Penicillium Roqueforti
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''Penicillium roqueforti'' is a common saprotrophic
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''
Penicillium ''Penicillium'' () is a genus of Ascomycota, ascomycetous fungus, fungi that is part of the mycobiome of many species and is of major importance in the natural environment, in food spoilage, and in food and drug production. Some members of th ...
''. 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 wat ...
s,
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...
s, and other
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
s. The fungus has been a constituent of
Roquefort Roquefort () is a sheep milk blue cheese from southern France. Though similar cheeses are produced elsewhere, European Union law, EU law dictates that only those cheeses aged in the natural Combalou caves of Roquefort-sur-Soulzon may bear the na ...
, Stilton, Danish blue, Cabrales, and other blue cheeses. Other blue cheeses, such as
Gorgonzola Gorgonzola (, ) is a famously pungent Italian blue cheese made from unskimmed cow's milk; believed to have been created in the 9th century; now with use of its name controlled under the criteria of a Protected Designation of Origin (PDO). Gorg ...
, are made with '' Penicillium glaucum''.


Classification

First described by the 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 (physical form and structure), its developmental processes, its biochemical and physiological propert ...
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 analysis of
ribosomal DNA The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
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 () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
. In 2014, researchers reported inducing the growth of sexual structures in ''P. roqueforti'', including ascogonia, cleistothecia, and
ascospore In fungi, an ascospore is the sexual spore formed inside an ascus—the sac-like cell that defines the division Ascomycota, the largest and most diverse Division (botany), division of fungi. After two parental cell nucleus, nuclei fuse, the ascu ...
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'' can tolerate cold temperatures, low oxygen levels, and both alkali and weaker acid preservatives which allows the fungi to thrive and be found in dairy environments, such as cheese. On the other hand, it also spoils refrigerated foods and meats, breads, and
silage Silage is fodder made from green foliage crops which have been preserved by fermentation (food), fermentation to the point of souring. It is fed to cattle, sheep and other ruminants. The fermentation and storage process is called ''ensilage'', ' ...
.


Uses

The chief industrial use of this species is the production of blue cheeses, such as its namesake
Roquefort Roquefort () is a sheep milk blue cheese from southern France. Though similar cheeses are produced elsewhere, European Union law, EU law dictates that only those cheeses aged in the natural Combalou caves of Roquefort-sur-Soulzon may bear the na ...
, Bleu de Bresse, Bleu du Vercors-Sassenage, Brebiblu, Cabrales, Cambozola (Blue Brie), Cashel Blue, Danish blue, Swedish Ädelost, Polish Rokpol made from cow's milk, Fourme d'Ambert, Fourme de Montbrison, Lanark Blue, Shropshire Blue, and Stilton, and some varieties of
Bleu d'Auvergne Bleu d'Auvergne () is a French blue cheese, named for its place of origin in the Auvergne (province), Auvergne region of south-central France. It is made from cow's milk, and is one of the cheeses granted the Appellation d'origine contrôlée fr ...
and
Gorgonzola Gorgonzola (, ) is a famously pungent Italian blue cheese made from unskimmed cow's milk; believed to have been created in the 9th century; now with use of its name controlled under the criteria of a Protected Designation of Origin (PDO). Gorg ...
. (Other blue cheeses, including Bleu de Gex and Rochebaron, use '' Penicillium glaucum''.) When placed into cream and aerated, ''P. roqueforti'' produces concentrated blue cheese flavoring, a type of enzyme-modified cheese. A similar flavoring can be produced using other sources of fat such as coconut oil. 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 pathogenic bacteria, bacterial infections, and antibiotic medications are widely used in the therapy ...
s, flavours, and fragrances, uses not regulated under the U.S. Toxic Substances Control Act. Its texture is
chitin Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...
ous.


Secondary metabolites

Considerable evidence indicates that most strains are capable of producing harmful secondary metabolites (
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
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 In organic chemistry, an imine ( or ) is a functional group or organic compound containing a carbon–nitrogen double bond (). The nitrogen atom can be attached to a hydrogen or an organic group (R). The carbon atom has two additional single bon ...
.
Secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
s 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 nervous tissue, nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insult (medical), insultsSpencer 2000 that can adversely affect function ...
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 catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...
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 (whereas normal methane has the formula ). In formulas, the group is often abbreviated as ...
ketone In organic chemistry, a ketone is an organic compound with the structure , where R and R' can be a variety of carbon-containing substituents. Ketones contain a carbonyl group (a carbon-oxygen double bond C=O). The simplest ketone is acetone ( ...
s, including 2-heptanone. Recent research has shown significant differences in metabolite production between ''P. roqueforti'' populations. The cheese-making populations, particularly the non-Roquefort strains, produce fewer metabolites compared to non-cheese populations found in lumber and silage. The non-Roquefort population's inability to produce PR toxin stems from a
guanine Guanine () (symbol G or Gua) is one of the four main nucleotide bases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine ( uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside ...
to
adenine Adenine (, ) (nucleoside#List of nucleosides and corresponding nucleobases, symbol A or Ade) is a purine nucleotide base that is found in DNA, RNA, and Adenosine triphosphate, ATP. Usually a white crystalline subtance. The shape of adenine is ...
nuceltide substitution in ORF 11 of the PR toxin biosynthetic cluster, introducing a premature
stop codon In molecular biology, a stop codon (or termination codon) is a codon (nucleotide triplet within messenger RNA) that signals the termination of the translation process of the current protein. Most codons in messenger RNA correspond to the additio ...
. Similarly, these strains cannot produce mycophenolic acid due to a deletion in the
lipase In biochemistry, lipase ( ) refers to a class 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; howe ...
/ esterase domain of the ''mpaC'' gene. While Roquefort strains show no genetic mutations in PR toxin genes, they still do not produce the toxin, suggesting
downregulation In biochemistry, in the biology, biological context of organisms' regulation of gene expression and production of gene products, downregulation is the process by which a cell (biology), cell decreases the production and quantities of its cellular ...
of the pathway. The Termignon cheese population show intermediate metabolite profiles between cheese and non-cheese populations, producing low levels of PR toxin while showing the highest production of MPA-related compounds. Non-cheese populations maintain higher metabolite diversity, particularly in
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s and
terpenoid The terpenoids, also known as isoprenoids, are a class of naturally occurring organic compound, organic chemicals derived from the 5-carbon compound isoprene and its derivatives called terpenes, diterpenes, etc. While sometimes used interchangeabl ...
s, which may provide competitive advantages in more complex environments where fungi must compete with other microorganisms. The reduced toxin production in cheese strains likely results from either deliberate selection for safer strains during domestication or the degeneration of unused metabolic pathways in the cheese environment.


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 agency of the United States government tasked with environmental protection matters. President Richard Nixon proposed the establishment of EPA on July 9, 1970; it began operation on De ...
.'' {{Authority control roqueforti Molds used in food production Fungi described in 1906 Taxa named by Charles Thom Fungi in cultivation Fungus species