Penicillium: Difference between revisions

Penicillium
	Penicillium sp.
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Ascomycota
Class:	Eurotiomycetes
Order:	Eurotiales
Family:	Aspergillaceae
Genus:	Penicillium; Link (1809)
Type species
	Penicillium expansum; Link (1809)
Species
	over 300
Synonyms
	List Aspergillopsis Sopp (1912) ; Carpenteles Langeron (1922) ; Chromocleista Yaguchi & Udagawa (1993) ; Citromyces Wehmer (1893) ; Coremium Link (1809) ; Eladia G. Sm. (1961) ; Eupenicillium F. Ludw. (1892) ; Floccaria Grev. (1827); ; Hemicarpenteles A. K. Sarbhoy & Elphick (1968) ; Moniliger Letell. (1839) ; Pritzeliella Henn. (1903) ; Thysanophora W.B. Kendr. (1961) ; Toluromyces Delitsch (1943) ; Walzia Sorokin (1871) ;

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Penicillium (/ˌpɛnɪˈsɪliəm/) is a genus of ascomycetous 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 the genus produce penicillin, a molecule that is used as an antibiotic, which kills or stops the growth of certain kinds of bacteria. Other species are used in cheesemaking. According to the Dictionary of the Fungi (10th edition, 2008), the widespread genus contains over 300 species.^[2]

Taxonomy

The genus was first described in the scientific literature by Johann Heinrich Friedrich Link in his 1809 work Observationes in ordines plantarum naturales; he wrote, "Penicillium. Thallus e floccis caespitosis septatis simplicibus aut ramosis fertilibus erectis apice penicillatis", (transl. Penicillium. The thallus consists of grassy tufts septated with simple or erect fertile branches with a brush tip) where penicillatis means "having tufts of fine hair".^[3]^[4] 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.^[5]^{[full citation needed]}

In his 1979 monograph, John I. Pitt divided Penicillium into four subgenera based on conidiophore morphology and branching pattern: Aspergilloides, Biverticillium, Furcatum, and Penicillium.^[6] Species included in subgenus Biverticillium were later merged into Talaromyces.

Species

Some penicillium mold on mandarin oranges, probably *Penicillium digitatum*.

Selected species include:

Penicillium albocoremium
Penicillium aurantiogriseum, a grain contaminant
Penicillium bilaiae, an agricultural inoculant
Penicillium camemberti, used in the production of Camembert, Brie and Cambozola cheeses
Penicillium candidum, 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 penicillin
Penicillium claviforme
Penicillium commune
Penicillium crustosum
Penicillium digitatum, a Citrus pathogen
Penicillium echinulatum produces Mycophenolic acid
Penicillium expansum, a pathogen of apples and other fruit, produces patulin
Penicillium glabrum
Penicillium glaucum, a mold that is used in the making of some types of blue cheese, including Bleu de Gex, Rochebaron, and some varieties of Bleu d'Auvergne and Gorgonzola.
Penicillium imranianum
Penicillium italicum, a Citrus pathogen
Penicillium lacussarmientei
Penicillium lusitanum, isolated from marine habitat
Penicillium purpurogenum
Penicillium roqueforti, used in making Roquefort, Danish Blue cheese, English Blue Stilton cheese, Gorgonzola cheese, and Cambozola
Penicillium stoloniferum
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.^[7]

Characteristics

The thallus (mycelium) consists of highly branched networks of multinucleated, usually colourless hyphae, with each pair of cells separated by a septum. Conidiophores are at the end of each branch accompanied by green spherical constricted units called conidia. These propagules play a significant role in reproduction; conidia are the main dispersal strategy of these fungi.^[8]

Sexual reproduction involves the production of ascospores, commencing with the fusion of an archegonium and an antheridium, with sharing of nuclei. The irregularly distributed asci 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 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 Penicillium.^[9] 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.^[10] 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;^[11] and P. allii, garlic.^[12] 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.^[13]

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.^[14] 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%.^[15]

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.^[16]

Economic value

Core structure of penicillin

Griseofulvin

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 and Penicillium roqueforti are the molds on Camembert, Brie, Roquefort, and many other cheeses. Penicillium nalgiovense 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.^[17]^[18]

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 pectinases, lipase, 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.^[19]

The genus includes a wide variety of species molds that are the source molds of major antibiotics. Penicillin, a drug produced by P. chrysogenum (formerly P. notatum), was accidentally discovered by Alexander Fleming in 1929, and found to inhibit the growth of Gram-positive bacteria (see beta-lactams). Its potential as an antibiotic was realized in the late 1930s, and Howard Florey and Ernst Chain 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.^[20]

Griseofulvin is an antifungal drug and a potential chemotherapeutic agent^[21] that was discovered in P. griseofulvum.^[22] Additional species that produce compounds capable of inhibiting the growth of tumor cells in vitro include: P. pinophilum,^[23] P. canescens,^[24] and P. glabrum.^[24]

Reproduction

Although many eukaryotes 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 occurs even in some of the supposedly asexual species. For example, sexual capability was recently shown for the fungus Penicillium roqueforti, used as a starter for blue cheese production.^[25] 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.^[26] 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 eukaryotes.^[27] 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" 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

^ Page Penicillium on "Mycobank". Westerdijk Fungal Biodiversity Institute. Archived from the original on 2024-02-29. Retrieved 2023-09-20.
^ Ainsworth GC (2008). Ainsworth & Bisby's Dictionary of the Fungi. CABI. p. 505. ISBN 978-0-85199-826-8.
^ Visagie CM, Houbraken J, Frisvad JC, Hong SB, Klaassen CH, Perrone G, et al. (June 2014). "Identification and nomenclature of the genus Penicillium". Studies in Mycology. 78 (1): 343–371. doi:10.1016/j.simyco.2014.09.001. PMC 4261876. PMID 25505353.
^ Link JH (1809). Der Gesellschaft Naturforschender Freunde zu Berlin Magazin für die neuesten Entdeckungen in der gesammten Naturkunde (in Latin). Vol. 3. Realschulbuchhandlung. pp. 3–42.
^ Samson RA, Pitt JI, eds. (July 1985). "Advances in penicillium and aspergillus systematics". Advances in penicillium and aspergillus systematics. First International Penicillium and Aspergillus NATO Workshop. Trippenhuis of the Royal Dutch Academy of Sciences and Letters: Plenum Press. ISBN 978-0-306-42222-5 – via Internet Archive.
^ Pitt JI (1979). The Genus Penicillium and Its Teleomorphic States Eupenicillium and Talaromyces. Academic Press. ISBN 978-0-12-557750-2.
^ Haubrich WS (2003). Medical Meanings: A Glossary of Word Origins. ACP Press. p. 175. ISBN 978-1-930513-49-5.
^ Pitt J (1985). "A laboratory guide to common Penicillium species". Mycologia. 79: 491. doi:10.2307/3807483. JSTOR 3807483. S2CID 84610634.
^ Samson RA, Seifert KA, Kuijpers AF, Houbraken JA, Frisvad JC (2004). "Phylogenetic analysis of Penicillium subgenus Penicillium using partial β-tubulin sequences" (PDF). Studies in Mycology. 49: 175–200.
^ Pitt JI, Basílico JC, Abarca ML, López C (2000). "Mycotoxins and toxigenic fungi". Medical Mycology. 38 (Suppl 1): 41–46. doi:10.1080/mmy.38.s1.41.46. PMID 11204163.
^ Baigrie B (2003-04-02). Taints and Off-Flavours in Foods. Elsevier Science. p. 134. ISBN 978-1-85573-449-4.
^ Valdez JG, Makuch MA, Ordovini AF, Masuelli RW, Overy DP, Piccolo RJ (2006). "First report of Penicillium allii as a field pathogen of garlic ( Allium sativum )". Plant Pathology. 55 (4): 583. doi:10.1111/j.1365-3059.2006.01411.x. hdl:11336/147639. ISSN 0032-0862.
^ da Costa GL, de Moraes AM, de Oliveira PC (1998). "Pathogenic action of Penicillium species on mosquito vectors of human tropical diseases". Journal of Basic Microbiology. 38 (5–6): 337–341. doi:10.1002/(sici)1521-4028(199811)38:5/6<337::aid-jobm337>3.3.co;2-e. PMID 9871331.
^ Fairs A, Wardlaw AJ, Pashley CH (2010). "Guidelines on ambient intramural airborne fungal spores". Journal of Investigational Allergology & Clinical Immunology. 20 (6): 490–498. PMID 21243933.
^ Chang JC, Foarde KK, Vanosdell DW (1995-01-01). "Growth evaluation of fungi ( Penicillium and Aspergillus spp.) on ceiling tiles". Atmospheric Environment. 29 (17): 2331–2337. Bibcode:1995AtmEn..29.2331C. doi:10.1016/1352-2310(95)00062-4. ISSN 1352-2310.
^ Semenov SA, Gumargalieva KZ, Zaikov GE (2003-09-24). Biodegradation and Durability of Materials Under the Effect of Microorganisms. VSP. pp. 34–35. ISBN 978-90-6764-388-7.
^ Mrázek J, Pachlová V, Buňka F, Černíková M, Dráb V, Bejblová M, et al. (May 2016). "Effects of different strains Penicillium nalgiovense in the Nalžovy cheese during ripening". Journal of the Science of Food and Agriculture. 96 (7): 2547–2554. Bibcode:2016JSFA...96.2547M. doi:10.1002/jsfa.7375. PMID 26251231.
^ Marianski S, Mariański A (2009). The Art of Making Fermented Sausages. Bookmagic LLC. p. 47. ISBN 978-0-9824267-1-5.
^ Leitão AL (April 2009). "Potential of Penicillium species in the bioremediation field". International Journal of Environmental Research and Public Health. 6 (4): 1393–1417. doi:10.3390/ijerph6041393. PMC 2681198. PMID 19440525.
^ Rifkind D, Freeman G (2005-05-20). The Nobel Prize Winning Discoveries in Infectious Diseases. Elsevier. pp. 43–46. ISBN 978-0-08-045957-8.
^ Singh P, Rathinasamy K, Mohan R, Panda D (June 2008). "Microtubule assembly dynamics: an attractive target for anticancer drugs". IUBMB Life. 60 (6): 368–375. doi:10.1002/iub.42. PMID 18384115.
^ De Carli L, Larizza L (March 1988). "Griseofulvin". Mutation Research. 195 (2): 91–126. doi:10.1016/0165-1110(88)90020-6. PMID 3277037.
^ Nicoletti R, Manzo E, Ciavatta ML (March 2009). "Occurence and bioactivities of funicone-related compounds[sic]". International Journal of Molecular Sciences. 10 (4): 1430–1444. doi:10.3390/ijms10041430. PMC 2680625. PMID 19468317.
^ ^a ^b Nicoletti R, Buommino E, De Filippis A, Lopez-Gresa MP, Manzo E, Carella A, et al. (2008-02-01). "Bioprospecting for antagonistic Penicillium strains as a resource of new antitumor compounds". World Journal of Microbiology and Biotechnology. 24 (2): 189–195. doi:10.1007/s11274-007-9455-y. ISSN 1573-0972.
^ Ropars J, Dupont J, Fontanillas E, Rodríguez de la Vega RC, Malagnac F, Coton M, et al. (2012). "Sex in cheese: evidence for sexuality in the fungus Penicillium roqueforti". PLOS ONE. 7 (11): e49665. Bibcode:2012PLoSO...749665R. doi:10.1371/journal.pone.0049665. PMC 3504111. PMID 23185400.
^ Böhm J, Hoff B, O'Gorman CM, Wolfers S, Klix V, Binger D, et al. (January 2013). "Sexual reproduction and mating-type-mediated strain development in the penicillin-producing fungus Penicillium chrysogenum". Proceedings of the National Academy of Sciences of the United States of America. 110 (4): 1476–1481. doi:10.1073/pnas.1217943110. PMC 3557024. PMID 23307807.
^ Malik SB, Pightling AW, Stefaniak LM, Schurko AM, Logsdon JM (August 2007). "An expanded inventory of conserved meiotic genes provides evidence for sex in Trichomonas vaginalis". PLOS ONE. 3 (8): e2879. Bibcode:2008PLoSO...3.2879M. doi:10.1371/journal.pone.0002879. PMC 2488364. PMID 18663385.

External links

[MycobankPenicillium-1] Page Penicillium on "Mycobank". Westerdijk Fungal Biodiversity Institute. Archived from the original on 2024-02-29. Retrieved 2023-09-20.

[2] Ainsworth GC (2008). Ainsworth & Bisby's Dictionary of the Fungi. CABI. p. 505. ISBN 978-0-85199-826-8.

[3] Visagie CM, Houbraken J, Frisvad JC, Hong SB, Klaassen CH, Perrone G, et al. (June 2014). "Identification and nomenclature of the genus Penicillium". Studies in Mycology. 78 (1): 343–371. doi:10.1016/j.simyco.2014.09.001. PMC 4261876. PMID 25505353.

[4] Link JH (1809). Der Gesellschaft Naturforschender Freunde zu Berlin Magazin für die neuesten Entdeckungen in der gesammten Naturkunde (in Latin). Vol. 3. Realschulbuchhandlung. pp. 3–42.

[5] Samson RA, Pitt JI, eds. (July 1985). "Advances in penicillium and aspergillus systematics". Advances in penicillium and aspergillus systematics. First International Penicillium and Aspergillus NATO Workshop. Trippenhuis of the Royal Dutch Academy of Sciences and Letters: Plenum Press. ISBN 978-0-306-42222-5 – via Internet Archive.

[6] Pitt JI (1979). The Genus Penicillium and Its Teleomorphic States Eupenicillium and Talaromyces. Academic Press. ISBN 978-0-12-557750-2.

[7] Haubrich WS (2003). Medical Meanings: A Glossary of Word Origins. ACP Press. p. 175. ISBN 978-1-930513-49-5.

[8] Pitt J (1985). "A laboratory guide to common Penicillium species". Mycologia. 79: 491. doi:10.2307/3807483. JSTOR 3807483. S2CID 84610634.

[9] Samson RA, Seifert KA, Kuijpers AF, Houbraken JA, Frisvad JC (2004). "Phylogenetic analysis of Penicillium subgenus Penicillium using partial β-tubulin sequences" (PDF). Studies in Mycology. 49: 175–200.

[10] Pitt JI, Basílico JC, Abarca ML, López C (2000). "Mycotoxins and toxigenic fungi". Medical Mycology. 38 (Suppl 1): 41–46. doi:10.1080/mmy.38.s1.41.46. PMID 11204163.

[11] Baigrie B (2003-04-02). Taints and Off-Flavours in Foods. Elsevier Science. p. 134. ISBN 978-1-85573-449-4.

[12] Valdez JG, Makuch MA, Ordovini AF, Masuelli RW, Overy DP, Piccolo RJ (2006). "First report of Penicillium allii as a field pathogen of garlic ( Allium sativum )". Plant Pathology. 55 (4): 583. doi:10.1111/j.1365-3059.2006.01411.x. hdl:11336/147639. ISSN 0032-0862.

[13] Costa GL, de Moraes AM, de Oliveira PC (1998). "Pathogenic action of Penicillium species on mosquito vectors of human tropical diseases". Journal of Basic Microbiology. 38 (5–6): 337–341. doi:10.1002/(sici)1521-4028(199811)38:5/6<337::aid-jobm337>3.3.co;2-e. PMID 9871331.

[14] Fairs A, Wardlaw AJ, Pashley CH (2010). "Guidelines on ambient intramural airborne fungal spores". Journal of Investigational Allergology & Clinical Immunology. 20 (6): 490–498. PMID 21243933.

[15] Chang JC, Foarde KK, Vanosdell DW (1995-01-01). "Growth evaluation of fungi ( Penicillium and Aspergillus spp.) on ceiling tiles". Atmospheric Environment. 29 (17): 2331–2337. Bibcode:1995AtmEn..29.2331C. doi:10.1016/1352-2310(95)00062-4. ISSN 1352-2310.

[16] Semenov SA, Gumargalieva KZ, Zaikov GE (2003-09-24). Biodegradation and Durability of Materials Under the Effect of Microorganisms. VSP. pp. 34–35. ISBN 978-90-6764-388-7.

[17] Mrázek J, Pachlová V, Buňka F, Černíková M, Dráb V, Bejblová M, et al. (May 2016). "Effects of different strains Penicillium nalgiovense in the Nalžovy cheese during ripening". Journal of the Science of Food and Agriculture. 96 (7): 2547–2554. Bibcode:2016JSFA...96.2547M. doi:10.1002/jsfa.7375. PMID 26251231.

[18] Marianski S, Mariański A (2009). The Art of Making Fermented Sausages. Bookmagic LLC. p. 47. ISBN 978-0-9824267-1-5.

[19] Leitão AL (April 2009). "Potential of Penicillium species in the bioremediation field". International Journal of Environmental Research and Public Health. 6 (4): 1393–1417. doi:10.3390/ijerph6041393. PMC 2681198. PMID 19440525.

[20] Rifkind D, Freeman G (2005-05-20). The Nobel Prize Winning Discoveries in Infectious Diseases. Elsevier. pp. 43–46. ISBN 978-0-08-045957-8.

[21] Singh P, Rathinasamy K, Mohan R, Panda D (June 2008). "Microtubule assembly dynamics: an attractive target for anticancer drugs". IUBMB Life. 60 (6): 368–375. doi:10.1002/iub.42. PMID 18384115.

[22] De Carli L, Larizza L (March 1988). "Griseofulvin". Mutation Research. 195 (2): 91–126. doi:10.1016/0165-1110(88)90020-6. PMID 3277037.

[23] Nicoletti R, Manzo E, Ciavatta ML (March 2009). "Occurence and bioactivities of funicone-related compounds[sic]". International Journal of Molecular Sciences. 10 (4): 1430–1444. doi:10.3390/ijms10041430. PMC 2680625. PMID 19468317.

[:0-24] Nicoletti R, Buommino E, De Filippis A, Lopez-Gresa MP, Manzo E, Carella A, et al. (2008-02-01). "Bioprospecting for antagonistic Penicillium strains as a resource of new antitumor compounds". World Journal of Microbiology and Biotechnology. 24 (2): 189–195. doi:10.1007/s11274-007-9455-y. ISSN 1573-0972.

[pmid23185400-25] Ropars J, Dupont J, Fontanillas E, Rodríguez de la Vega RC, Malagnac F, Coton M, et al. (2012). "Sex in cheese: evidence for sexuality in the fungus Penicillium roqueforti". PLOS ONE. 7 (11): e49665. Bibcode:2012PLoSO...749665R. doi:10.1371/journal.pone.0049665. PMC 3504111. PMID 23185400.

[pmid23307807-26] Böhm J, Hoff B, O'Gorman CM, Wolfers S, Klix V, Binger D, et al. (January 2013). "Sexual reproduction and mating-type-mediated strain development in the penicillin-producing fungus Penicillium chrysogenum". Proceedings of the National Academy of Sciences of the United States of America. 110 (4): 1476–1481. doi:10.1073/pnas.1217943110. PMC 3557024. PMID 23307807.

[pmid18663385-27] Malik SB, Pightling AW, Stefaniak LM, Schurko AM, Logsdon JM (August 2007). "An expanded inventory of conserved meiotic genes provides evidence for sex in Trichomonas vaginalis". PLOS ONE. 3 (8): e2879. Bibcode:2008PLoSO...3.2879M. doi:10.1371/journal.pone.0002879. PMC 2488364. PMID 18663385.

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@@ Line 1: / Line 1: @@
 {{Short description|Genus of fungi}}
-{{pp-pc1}}
 {{technical|date=August 2018}}
+{{cs1 config |name-list-style=vanc |display-authors=6}}
 {{Automatic taxobox
 | image = Penicillium Pengo.jpg
@@ Line 10: / Line 10: @@
 | type_species_authority = Link (1809)
 | subdivision_ranks = Species
-| subdivision = over 300<br/>
+| subdivision = [[List of Penicillium species|over 300]]
+| synonyms_ref = <ref name=MycobankPenicillium>Page ''Penicillium'' on {{cite web | url=https://www.mycobank.org/page/Name%20details%20page/39268 | title=Mycobank | publisher=[[Westerdijk Fungal Biodiversity Institute]] | access-date=2023-09-20 | archive-date=2024-02-29 | archive-url=https://web.archive.org/web/20240229055024/https://www.mycobank.org/page/Name%20details%20page/39268 | url-status=live }}</ref>
-[[List of Penicillium species]]
+| synonyms = {{Collapsible list|bullets=on
-| synonyms_ref = <ref name="urlMycoBank: Penicillium"/>
-| synonyms =''Floccaria'' <small>[[Grev.]] (1827)</small><br/>
+|''Aspergillopsis'' <small>Sopp (1912)</small>
-''Aspergilloides'' <small>[[R. P. Dierckx|Dierckx]] (1901)</small><br/>
+|''Carpenteles'' <small>Langeron (1922)</small>
-''Walzia'' <small>[[Nikolai Vasilyevich Sorokin|Sorokin]] (1871)</small><br/>
+|''Chromocleista'' <small>Yaguchi & Udagawa (1993)</small>
-''Pritzeliella'' <small>[[Henn.]] (1903)</small>
+|''Citromyces'' <small>Wehmer (1893)</small>
+|''Coremium'' <small>Link (1809)</small>
+|''Eladia'' <small>G. Sm. (1961)</small>
+|''Eupenicillium'' <small>F. Ludw. (1892)</small>
+|''Floccaria'' <small>[[Grev.]] (1827)</small><br/>
+|''Hemicarpenteles'' <small>A. K. Sarbhoy & Elphick (1968)</small>
+|''Moniliger'' <small>Letell. (1839)</small>
+|''Pritzeliella'' <small>[[Henn.]] (1903)</small>
+|''Thysanophora'' <small>W.B. Kendr. (1961)</small>
+|''Toluromyces'' <small>Delitsch (1943)</small>
+|''Walzia'' <small>[[Nikolai Vasilyevich Sorokin|Sorokin]] (1871)</small>
+}}
 }}
 '''''Penicillium''''' ({{IPAc-en|ˌ|p|ɛ|n|ɪ|ˈ|s|ɪ|l|i|əm}}) 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 the genus produce [[penicillin]], a molecule that is used as an [[antibiotic]], which kills or stops the growth of certain kinds of bacteria. Other species are used in [[cheesemaking]]. According to the ''Dictionary of the Fungi'' (10th edition, 2008), the widespread genus contains over 300 species.<ref name=Kirk2008/>
+Some members of the genus produce [[penicillin]], a molecule that is used as an [[antibiotic]], which kills or stops the growth of certain kinds of bacteria. Other species are used in [[cheesemaking]]. According to the ''Dictionary of the Fungi'' (10th edition, 2008), the widespread genus contains over 300 species.<ref>{{Cite book | vauthors = Ainsworth GC |url=https://books.google.com/books?id=IFD4_VFRDdUC |title=Ainsworth & Bisby's Dictionary of the Fungi |date=2008 |publisher=CABI |isbn=978-0-85199-826-8 |pages=505 |language=en}}</ref>
 ==Taxonomy==
-The genus was first described in the scientific literature by [[Johann Heinrich Friedrich Link]] 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".<ref>Identification and nomenclature of the genus Penicillium, C.M. Visagie1, J. Houbraken1, , , J.C. Frisvad2, , , S.-B. Hong3, C.H.W. Klaassen4, G. Perrone5, K.A. Seifert6, J. Varga7, T. Yaguchi8, R.A. Samson, 22 September 2014, https://dx.doi.org/10.1016/j.simyco.2014.09.001</ref><ref name=Link1809/> Link included three species—''[[Penicillium candidum|P.&nbsp;candidum]]'', ''[[Penicillium expansum|P.&nbsp;expansum]]'', and ''[[Penicillium glaucum|P.&nbsp;glaucum]]''—all of which produced a brush-like [[conidiophore]] (asexual spore-producing structure). The common apple rot fungus ''P.&nbsp;expansum'' was later selected as the [[type species]].<ref name=Samson1985/>
+The genus was first described in the scientific literature by [[Johann Heinrich Friedrich Link]] in his 1809 work {{lang|la|Observationes in ordines plantarum naturales}}; he wrote, {{lang|la|"Penicillium. Thallus e floccis caespitosis septatis simplicibus aut ramosis fertilibus erectis apice penicillatis"}}, ({{trans|Penicillium. The thallus consists of grassy tufts septated with simple or erect fertile branches with a brush tip}}) where {{lang|la|penicillatis}} means "having tufts of fine hair".<ref>{{cite journal | vauthors = Visagie CM, Houbraken J, Frisvad JC, Hong SB, Klaassen CH, Perrone G, Seifert KA, Varga J, Yaguchi T, Samson RA | title = Identification and nomenclature of the genus Penicillium | journal = Studies in Mycology | volume = 78 | issue = 1 | pages = 343–371 | date = June 2014 | pmid = 25505353 | pmc = 4261876 | doi = 10.1016/j.simyco.2014.09.001 }}</ref><ref>{{Cite book | vauthors = Link JH |url=https://gdz.sub.uni-goettingen.de/id/PPN608227714_0003?tify=%7B%22pages%22:%5B31%5D%7D |title=Der Gesellschaft Naturforschender Freunde zu Berlin Magazin für die neuesten Entdeckungen in der gesammten Naturkunde |date=1809 |publisher=Realschulbuchhandlung |edition= |volume=3 |pages=3–42 |language=la}}</ref> Link included three species—''[[Penicillium candidum|P.&nbsp;candidum]]'', ''[[Penicillium expansum|P.&nbsp;expansum]]'', and ''[[Penicillium glaucum|P.&nbsp;glaucum]]''—all of which produced a brush-like [[conidiophore]] (asexual spore-producing structure). The common apple rot fungus ''P.&nbsp;expansum'' was later selected as the [[type species]].<ref>{{Cite conference |book-title=Advances in penicillium and aspergillus systematics |editor1-first=Robert A. |editor1-last=Samson |editor2-first=John I. |editor2-last=Pitt |conference=First International Penicillium and Aspergillus NATO Workshop |location=Trippenhuis of the Royal Dutch Academy of Sciences and Letters |url=http://archive.org/details/advancesinpenici0000inte |url-access=registration |title=Advances in penicillium and aspergillus systematics |date=July 1985 |publisher=Plenum Press |via=Internet Archive |isbn=978-0-306-42222-5}}</ref>{{fcn|reason=article author not given|date=September 2024}}
-In his 1979 [[monograph]], John I. Pitt divided ''Penicillium'' into four [[subgenera]] based on conidiophore [[morphology (biology)|morphology]] and branching pattern: ''Aspergilloides'', ''Biverticillium'', ''Furcatum'', and ''Penicillium''.<ref name=Pitt1979/> Species included in subgenus ''Biverticillium'' were later merged into ''[[Talaromyces]]''.
+In his 1979 [[monograph]], John I. Pitt divided ''Penicillium'' into four [[subgenera]] based on conidiophore [[morphology (biology)|morphology]] and branching pattern: ''Aspergilloides'', ''Biverticillium'', ''Furcatum'', and ''Penicillium''.<ref>{{Cite book | vauthors = Pitt JI |url=https://books.google.com/books?id=W38yAAAAIAAJ |title=The Genus Penicillium and Its Teleomorphic States Eupenicillium and Talaromyces |date=1979 |publisher=Academic Press |isbn=978-0-12-557750-2 |language=en}}</ref> Species included in subgenus ''Biverticillium'' were later merged into ''[[Talaromyces]]''.
 === Species ===
@@ Line 33: / Line 44: @@
 [[File:Penicilliummandarijntjes.jpg|thumb|right|Some penicillium mold on [[mandarin orange]]s, probably ''Penicillium digitatum''.]]
-Selected species include;
+Selected species include:
 * ''[[Penicillium albocoremium]]''
 * ''[[Penicillium aurantiogriseum]]'', a grain contaminant
-* ''[[Penicillium bilaiae]]'', which is an agricultural inoculant
+* ''[[Penicillium bilaiae]]'', an agricultural inoculant
-* ''[[Penicillium camemberti]]'', which is used in the production of [[Camembert (cheese)|Camembert]] and [[Brie cheese|Brie]] [[cheese]]s
+* ''[[Penicillium camemberti]]'', used in the production of [[Camembert (cheese)|Camembert]], [[Brie cheese|Brie]] and [[Cambozola]] cheeses
 * ''[[Penicillium candidum]]'', 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]] [[penicillin]]
+* ''[[Penicillium chrysogenum]]'' (previously known as ''[[Penicillium notatum]]''), which produces the antibiotic [[penicillin]]
 * ''[[Penicillium claviforme]]''
 * ''[[Penicillium commune]]''
@@ Line 45: / Line 56: @@
 * ''[[Penicillium digitatum]]'', a ''Citrus'' pathogen
 * ''[[Penicillium echinulatum]]'' produces [[Mycophenolic acid]]
-* ''[[Penicillium expansum]]'', a pathogen of apples and other fruit, produces patulin
+* ''[[Penicillium expansum]]'', a pathogen of apples and other fruit, produces [[patulin]]
 * ''[[Penicillium glabrum]]''
+* ''[[Penicillium glaucum]]'', a [[Mold (fungus)|mold]] that is used in the making of some types of [[blue cheese]], including [[Bleu de Gex]], [[Rochebaron]], and some varieties of [[Bleu d'Auvergne]] and [[Gorgonzola cheese|Gorgonzola]].
 * ''[[Penicillium imranianum]]''
 * ''[[Penicillium italicum]]'', a ''Citrus'' pathogen
@@ Line 52: / Line 64: @@
 * ''[[Penicillium lusitanum]]'', isolated from marine habitat
 * ''[[Penicillium purpurogenum]]''
-* ''[[Penicillium roqueforti]]'', which is used in making [[Roquefort cheese|Roquefort]], [[Danish Blue cheese]], English Blue [[Stilton cheese]], and [[Gorgonzola cheese]]
+* ''[[Penicillium roqueforti]]'', used in making [[Roquefort cheese|Roquefort]], [[Danish Blue cheese]], English Blue [[Stilton cheese]], [[Gorgonzola cheese]], and [[Cambozola]]
 * ''[[Penicillium stoloniferum]]''
 * ''[[Penicillium ulaiense]]'', a ''Citrus'' pathogen in Asia
@@ Line 59: / Line 71: @@
 === 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.<ref name=Haubrich2003/>
+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.<ref>{{Cite book | vauthors = Haubrich WS |url=https://books.google.com/books?id=NXmlIwkQBLAC&pg=PA175 |title=Medical Meanings: A Glossary of Word Origins |date=2003 |publisher=ACP Press |isbn=978-1-930513-49-5 |pages=175 |language=en}}</ref>
 == Characteristics ==
 [[File:Penicillium Spp..jpg|thumb|''Penicillium'' sp. under bright field microscopy (10 × 100 magnification) with lactophenol cotton blue stain]]
-The [[thallus]] ([[mycelium]]) consists of highly branched networks of [[multinucleated]], usually colourless [[hyphae]], with each pair of cells separated by a [[septum]]. [[Conidiophores]] are at the end of each branch accompanied by green spherical constricted units called [[conidia]]. These propagules play a significant role in reproduction; conidia are the main dispersal strategy of these fungi.<ref>{{Cite journal|last=Pitt|first=J.|date=1985|title=A laboratory guide to common Penicillium species|url=https://www.semanticscholar.org/paper/A-laboratory-guide-to-common-Penicillium-species-Pitt/7c86b2f5283c16d926fa93de64ef535040eec860?p2df| volume=79 | doi=10.2307/3807483|jstor=3807483|s2cid=84610634}}</ref>
+The [[thallus]] ([[mycelium]]) consists of highly branched networks of [[multinucleated]], usually colourless [[hyphae]], with each pair of cells separated by a [[septum]]. [[Conidiophores]] are at the end of each branch accompanied by green spherical constricted units called [[conidia]]. These propagules play a significant role in reproduction; conidia are the main dispersal strategy of these fungi.<ref>{{Cite journal | vauthors = Pitt J |date=1985 |title=A laboratory guide to common Penicillium species |journal=Mycologia |volume=79 |page=491 |doi=10.2307/3807483 |jstor=3807483 |s2cid=84610634}}</ref>
 Sexual reproduction involves the production of [[ascospore]]s, commencing with the fusion of an [[archegonium]] and an [[antheridium]], with sharing of nuclei. The irregularly distributed [[ascus|asci]] 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]] and live mainly on organic biodegradable substances. Commonly known in America as [[Mold (fungus)|mold]]s, they are among the main causes of [[food spoilage]], especially species of [[subgenus]] ''Penicillium''.<ref name=Samson2004/> Many species produce highly toxic [[mycotoxin]]s. 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.<ref name=Pitt2000/> Some species have a blue color, commonly growing on old bread and giving it a blue fuzzy texture.
+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]] and live mainly on organic biodegradable substances. Commonly known in America as [[Mold (fungus)|mold]]s, they are among the main causes of [[food spoilage]], especially species of [[subgenus]] ''Penicillium''.<ref>{{Cite journal | vauthors = Samson RA, Seifert KA, Kuijpers AF, Houbraken JA, Frisvad JC |date=2004 |title=Phylogenetic analysis of Penicillium subgenus Penicillium using partial β-tubulin sequences |url=https://www.studiesinmycology.org/sim/Sim49/Sim49Article2.pdf |journal=Studies in Mycology |volume=49 |pages=175–200}}</ref> Many species produce highly toxic [[mycotoxin]]s. 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.<ref>{{cite journal | vauthors = Pitt JI, Basílico JC, Abarca ML, López C | title = Mycotoxins and toxigenic fungi | journal = Medical Mycology | volume = 38 | pages = 41–46 | date = 2000 | issue = Suppl 1 | doi = 10.1080/mmy.38.s1.41.46 | pmid = 11204163 }}</ref> 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 ''[[Penicillium expansum|P.&nbsp;expansum]]'', apples and pears; ''[[Penicillium digitatum|P.&nbsp;digitatum]]'', citrus fruits;<ref name=Baigre2003/> and ''[[Penicillium allii|P.&nbsp;allii]]'', garlic.<ref name=Valdez2006/> Some species are known to be pathogenic to animals; ''[[Penicillium corylophilum|P.&nbsp;corylophilum]]'', ''[[Penicillium fellutanum|P.&nbsp;fellutanum]]'', ''[[Penicillium implicatum|P.&nbsp;implicatum]]'', ''[[Penicillium janthinellum|P.&nbsp;janthinellum]]'', ''[[Penicillium viridicatum|P.&nbsp;viridicatum]]'', and ''[[Penicillium waksmanii|P.&nbsp;waksmanii]]'' are potential pathogens of [[mosquitoes]].<ref name=daCosta1998/>
+Some ''Penicillium'' species affect the fruits and bulbs of plants, including ''[[Penicillium expansum|P.&nbsp;expansum]]'', apples and pears; ''[[Penicillium digitatum|P.&nbsp;digitatum]]'', citrus fruits;<ref>{{Cite book | vauthors = Baigrie B  |url=https://books.google.com/books?id=xDcNCY31oQ4C&pg=PA134 |title=Taints and Off-Flavours in Foods |date=2003-04-02 |publisher=Elsevier Science |isbn=978-1-85573-449-4 |pages=134 |language=en}}</ref> and ''[[Penicillium allii|P.&nbsp;allii]]'', garlic.<ref>{{Cite journal | vauthors = Valdez JG, Makuch MA, Ordovini AF, Masuelli RW, Overy DP, Piccolo RJ |date=2006 |title=First report of Penicillium allii as a field pathogen of garlic ( Allium sativum ) |journal=Plant Pathology |language=en |volume=55 |issue=4 |pages=583 |doi=10.1111/j.1365-3059.2006.01411.x |issn=0032-0862|hdl=11336/147639 |hdl-access=free }}</ref> Some species are known to be pathogenic to animals; ''[[Penicillium corylophilum|P.&nbsp;corylophilum]]'', ''[[Penicillium fellutanum|P.&nbsp;fellutanum]]'', ''[[Penicillium implicatum|P.&nbsp;implicatum]]'', ''[[Penicillium janthinellum|P.&nbsp;janthinellum]]'', ''[[Penicillium viridicatum|P.&nbsp;viridicatum]]'', and ''[[Penicillium waksmanii|P.&nbsp;waksmanii]]'' are potential pathogens of [[mosquitoes]].<ref>{{cite journal | vauthors = da Costa GL, de Moraes AM, de Oliveira PC | title = Pathogenic action of Penicillium species on mosquito vectors of human tropical diseases | journal = Journal of Basic Microbiology | volume = 38 | issue = 5–6 | pages = 337–341 | date = 1998 | pmid = 9871331 | doi = 10.1002/(sici)1521-4028(199811)38:5/6<337::aid-jobm337>3.3.co;2-e }}</ref>
-''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.<ref name=Fairs2010/> Even [[Tile#Ceiling tiles|ceiling tile]]s 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%.<ref name=Chang1995/>
+''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.<ref>{{cite journal | vauthors = Fairs A, Wardlaw AJ, Pashley CH | title = Guidelines on ambient intramural airborne fungal spores | journal = Journal of Investigational Allergology & Clinical Immunology | volume = 20 | issue = 6 | pages = 490–498 | date = 2010 | pmid = 21243933 }}</ref> Even [[Tile#Ceiling tiles|ceiling tile]]s 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%.<ref>{{Cite journal | vauthors = Chang JC, Foarde KK, Vanosdell DW |date=1995-01-01 |title=Growth evaluation of fungi ( Penicillium and Aspergillus spp.) on ceiling tiles |url=https://ui.adsabs.harvard.edu/abs/1995AtmEn..29.2331C |journal=Atmospheric Environment |volume=29 |issue=17 |pages=2331–2337 |doi=10.1016/1352-2310(95)00062-4 |bibcode=1995AtmEn..29.2331C |issn=1352-2310}}</ref>
-Some ''Penicillium'' species cause damage to machinery and the combustible materials and lubricants used to run and maintain them. For example, ''[[Penicillium chrysogenum|P.&nbsp;chrysogenum]] (formerly [[P. chrysogenum|P. notatum]])'', ''[[Penicillium steckii|P.&nbsp;steckii]]'', ''[[Penicillium notatum|&nbsp;]][[Penicillium cyclopium|P.&nbsp;cyclopium]]'', and ''[[Penicillium nalgiovensis|P.&nbsp;nalgiovensis]]'' affect fuels; ''P.&nbsp;chrysogenum'', ''[[Penicillium rubrum|P.&nbsp;rubrum]]'', and ''[[Penicillium verrucosum|P.&nbsp;verrucosum]]'' cause damage to oils and lubricants; ''[[Penicillium regulosum|P.&nbsp;regulosum]]'' damages optical and protective glass.<ref name=Semenov2003/>
+Some ''Penicillium'' species cause damage to machinery and the combustible materials and lubricants used to run and maintain them. For example, ''[[Penicillium chrysogenum|P.&nbsp;chrysogenum]] (formerly [[P. chrysogenum|P. notatum]])'', ''[[Penicillium steckii|P.&nbsp;steckii]]'', ''[[Penicillium notatum|&nbsp;]][[Penicillium cyclopium|P.&nbsp;cyclopium]]'', and ''[[Penicillium nalgiovensis|P.&nbsp;nalgiovensis]]'' affect fuels; ''P.&nbsp;chrysogenum'', ''[[Penicillium rubrum|P.&nbsp;rubrum]]'', and ''[[Penicillium verrucosum|P.&nbsp;verrucosum]]'' cause damage to oils and lubricants; ''[[Penicillium regulosum|P.&nbsp;regulosum]]'' damages optical and protective glass.<ref>{{Cite book | vauthors = Semenov SA, Gumargalieva KZ, Zaikov GE |url=https://books.google.com/books?id=Kj89NRHHqPMC&pg=PA35 |title=Biodegradation and Durability of Materials Under the Effect of Microorganisms  |date=2003-09-24 |publisher=VSP |isbn=978-90-6764-388-7 |pages=34–35 |language=en}}</ref>
 == Economic value ==
@@ Line 87: / Line 99: @@
 | caption2  = Griseofulvin
 }}
-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]]'' and ''[[Penicillium roqueforti]]'' are the molds on [[Camembert]], [[Brie]], [[Roquefort]], and many other cheeses. ''[[Penicillium nalgiovense]]'' 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.<ref name="Mrázek2015"/><ref name=Marianski2009/>
+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]]'' and ''[[Penicillium roqueforti]]'' are the molds on [[Camembert]], [[Brie]], [[Roquefort]], and many other cheeses. ''[[Penicillium nalgiovense]]'' 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.<ref>{{cite journal | vauthors = Mrázek J, Pachlová V, Buňka F, Černíková M, Dráb V, Bejblová M, Staněk K, Buňková L | title = Effects of different strains Penicillium nalgiovense in the Nalžovy cheese during ripening | journal = Journal of the Science of Food and Agriculture | volume = 96 | issue = 7 | pages = 2547–2554 | date = May 2016 | pmid = 26251231 | doi = 10.1002/jsfa.7375 | bibcode = 2016JSFA...96.2547M }}</ref><ref>{{Cite book | vauthors = Marianski S, Mariański A  |url=https://books.google.com/books?id=3uzSvCdVqYwC&pg=PA47 |title=The Art of Making Fermented Sausages |date=2009 |publisher=Bookmagic LLC |isbn=978-0-9824267-1-5 |pages=47 |language=en}}</ref>
-In addition to their importance in the food industry, species of ''Penicillium'' and ''Aspergillus'' serve in the production of a number of biotechnologically produced [[enzyme]]s and other macromolecules, such as [[gluconic acid|gluconic]], [[citric acid|citric]], and [[tartaric acid]]s, as well as several [[pectinase]]s, [[lipase]], [[amylase]]s, [[cellulase]]s, and [[protease]]s.  Some ''Penicillium'' species have shown potential for use in [[bioremediation]], more specifically [[mycoremediation]], because of their ability to break down a variety of [[Environmental xenobiotic|xenobiotic compounds]].<ref name=Leitao2009/>
+In addition to their importance in the food industry, species of ''Penicillium'' and ''Aspergillus'' serve in the production of a number of biotechnologically produced [[enzyme]]s and other macromolecules, such as [[gluconic acid|gluconic]], [[citric acid|citric]], and [[tartaric acid]]s, as well as several [[pectinase]]s, [[lipase]], [[amylase]]s, [[cellulase]]s, and [[protease]]s. Some ''Penicillium'' species have shown potential for use in [[bioremediation]], more specifically [[mycoremediation]], because of their ability to break down a variety of [[Environmental xenobiotic|xenobiotic compounds]].<ref>{{cite journal | vauthors = Leitão AL | title = Potential of Penicillium species in the bioremediation field | journal = International Journal of Environmental Research and Public Health | volume = 6 | issue = 4 | pages = 1393–1417 | date = April 2009 | pmid = 19440525 | pmc = 2681198 | doi = 10.3390/ijerph6041393 | doi-access = free }}</ref>
-The genus includes a wide variety of species molds that are the source molds of major [[antibiotic]]s. [[Penicillin]], a drug produced by ''[[Penicillium rubens|P.&nbsp;rubens]]'', was accidentally discovered by [[Alexander Fleming]] in 1929, and found to inhibit the growth of [[Gram-positive]] bacteria (see [[beta-lactams]]). Its potential as an antibiotic was realized in the late 1930s, and [[Howard Florey]] and [[Ernst Chain]] 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.<ref name=Rifkind2005/>
+The genus includes a wide variety of species molds that are the source molds of major [[antibiotic]]s. [[Penicillin]], a drug produced by ''[[Penicillium chrysogenum|P.&nbsp;chrysogenum]] (formerly P.&nbsp;notatum),'' was accidentally discovered by [[Alexander Fleming]] in 1929, and found to inhibit the growth of [[Gram-positive]] bacteria (see [[beta-lactams]]). Its potential as an antibiotic was realized in the late 1930s, and [[Howard Florey]] and [[Ernst Chain]] 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.<ref>{{Cite book | vauthors = Rifkind D, Freeman G |url=https://books.google.com/books?id=d3wdy3b9VUkC&pg=PA43 |title=The Nobel Prize Winning Discoveries in Infectious Diseases |date=2005-05-20 |publisher=Elsevier |isbn=978-0-08-045957-8 |pages=43–46 |language=en}}</ref>
-[[Griseofulvin]] is an [[antifungal drug]] and a potential [[chemotherapeutic agent]]<ref name=Singh2008/> that was discovered in ''[[Penicillium griseofulvum|P.&nbsp;griseofulvum]]''.<ref name=DeCarli1988/> Additional species that produce compounds capable of inhibiting the growth of tumor cells ''[[in vitro]]'' include: ''[[Penicillium pinophilum|P.&nbsp;pinophilum]]'',<ref name=Nicoletti2009/> ''[[Penicillium canescens|P.&nbsp;canescens]]'',<ref name=Nicoletti2008/> and ''[[Penicillium glabrum|P.&nbsp;glabrum]]''.<ref name=Nicoletti2008/>
+[[Griseofulvin]] is an [[antifungal drug]] and a potential [[chemotherapeutic agent]]<ref>{{cite journal | vauthors = Singh P, Rathinasamy K, Mohan R, Panda D | title = Microtubule assembly dynamics: an attractive target for anticancer drugs | journal = IUBMB Life | volume = 60 | issue = 6 | pages = 368–375 | date = June 2008 | pmid = 18384115 | doi = 10.1002/iub.42 }}</ref> that was discovered in ''[[Penicillium griseofulvum|P.&nbsp;griseofulvum]]''.<ref>{{cite journal | vauthors = De Carli L, Larizza L | title = Griseofulvin | journal = Mutation Research | volume = 195 | issue = 2 | pages = 91–126 | date = March 1988 | pmid = 3277037 | doi = 10.1016/0165-1110(88)90020-6 }}</ref> Additional species that produce compounds capable of inhibiting the growth of tumor cells ''[[in vitro]]'' include: ''[[Penicillium pinophilum|P.&nbsp;pinophilum]]'',<ref>{{cite journal |vauthors=Nicoletti R, Manzo E, Ciavatta ML |date=March 2009 |title=Occurence and bioactivities of funicone-related compounds[sic] |journal=International Journal of Molecular Sciences |volume=10 |issue=4 |pages=1430–1444 |doi=10.3390/ijms10041430 |pmc=2680625 |pmid=19468317 |doi-access=free}}</ref> ''[[Penicillium canescens|P.&nbsp;canescens]]'',<ref name=":0">{{Cite journal | vauthors = Nicoletti R, Buommino E, De Filippis A, Lopez-Gresa MP, Manzo E, Carella A, Petrazzuolo M, Tufano MA |date=2008-02-01 |title=Bioprospecting for antagonistic Penicillium strains as a resource of new antitumor compounds |journal=World Journal of Microbiology and Biotechnology |language=en |volume=24 |issue=2 |pages=189–195 |doi=10.1007/s11274-007-9455-y |issn=1573-0972}}</ref> and ''[[Penicillium glabrum|P.&nbsp;glabrum]]''.<ref name=":0" />
 ==Reproduction==
-Although many [[eukaryote]]s are able to reproduce [[Sexual reproduction|sexually]], as much as 20% of [[fungus|fungal]] species had been thought to reproduce exclusively by [[Asexual reproduction|asexual]] means. However recent studies have revealed that [[sex]] occurs even in some of the supposedly asexual species.  For example, sexual capability was recently shown for the fungus ''[[Penicillium roqueforti]]'', used as a starter for [[blue cheese]] production.<ref name="pmid23185400">{{cite journal |vauthors=Ropars J, Dupont J, Fontanillas E, Rodríguez de la Vega RC, Malagnac F, Coton M, Giraud T, López-Villavicencio M |title=Sex in cheese: evidence for sexuality in the fungus Penicillium roqueforti |journal=PLOS ONE |volume=7 |issue=11 |pages=e49665 |year=2012 |pmid=23185400 |pmc=3504111 |doi=10.1371/journal.pone.0049665 |bibcode=2012PLoSO...749665R |doi-access=free }}</ref>  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.<ref name="pmid23307807">{{cite journal |vauthors=Böhm J, Hoff B, O'Gorman CM, Wolfers S, Klix V, Binger D, Zadra I, Kürnsteiner H, Pöggeler S, Dyer PS, Kück U |title=Sexual reproduction and mating-type-mediated strain development in the penicillin-producing fungus Penicillium chrysogenum |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=110 |issue=4 |pages=1476–81 |date=January 2013  |pmid=23307807 |pmc=3557024 |doi=10.1073/pnas.1217943110 |doi-access=free }}</ref> finally demonstrated sexual reproduction in ''P. chrysogenum''.
+Although many [[eukaryote]]s are able to reproduce [[Sexual reproduction|sexually]], as much as 20% of [[fungus|fungal]] species had been thought to reproduce exclusively by [[Asexual reproduction|asexual]] means. However recent studies have revealed that [[sex]] occurs even in some of the supposedly asexual species. For example, sexual capability was recently shown for the fungus ''[[Penicillium roqueforti]]'', used as a starter for [[blue cheese]] production.<ref name="pmid23185400">{{cite journal | vauthors = Ropars J, Dupont J, Fontanillas E, Rodríguez de la Vega RC, Malagnac F, Coton M, Giraud T, López-Villavicencio M | title = Sex in cheese: evidence for sexuality in the fungus Penicillium roqueforti | journal = PLOS ONE | volume = 7 | issue = 11 | pages = e49665 | year = 2012 | pmid = 23185400 | pmc = 3504111 | doi = 10.1371/journal.pone.0049665 | doi-access = free | bibcode = 2012PLoSO...749665R }}</ref> 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.<ref name="pmid23307807">{{cite journal | vauthors = Böhm J, Hoff B, O'Gorman CM, Wolfers S, Klix V, Binger D, Zadra I, Kürnsteiner H, Pöggeler S, Dyer PS, Kück U | title = Sexual reproduction and mating-type-mediated strain development in the penicillin-producing fungus Penicillium chrysogenum | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 110 | issue = 4 | pages = 1476–1481 | date = January 2013 | pmid = 23307807 | pmc = 3557024 | doi = 10.1073/pnas.1217943110 | doi-access = free }}</ref> finally demonstrated sexual reproduction in ''P. chrysogenum''.
-These findings with ''Penicillium'' species are consistent with accumulating evidence from studies of other [[Eukaryote|eukaryotic]] species that sex was likely present in the common ancestor of all [[eukaryote]]s.<ref name="pmid18663385">{{cite journal |vauthors=Malik SB, Pightling AW, Stefaniak LM, Schurko AM, Logsdon JM |title=An expanded inventory of conserved meiotic genes provides evidence for sex in Trichomonas vaginalis |journal=PLOS ONE |volume=3 |issue=8 |pages=e2879 |year=2008 |pmid=18663385 |pmc=2488364 |doi=10.1371/journal.pone.0002879 |bibcode=2008PLoSO...3.2879M |doi-access=free }}</ref>  Furthermore, these recent results suggest that sex can be maintained even when very little [[genetic variability]] is produced.
+These findings with ''Penicillium'' species are consistent with accumulating evidence from studies of other [[Eukaryote|eukaryotic]] species that sex was likely present in the common ancestor of all [[eukaryote]]s.<ref name="pmid18663385">{{cite journal | vauthors = Malik SB, Pightling AW, Stefaniak LM, Schurko AM, Logsdon JM | title = An expanded inventory of conserved meiotic genes provides evidence for sex in Trichomonas vaginalis | journal = PLOS ONE | volume = 3 | issue = 8 | pages = e2879 | date = August 2007 | pmid = 18663385 | pmc = 2488364 | doi = 10.1371/journal.pone.0002879 | doi-access = free | bibcode = 2008PLoSO...3.2879M }}</ref> Furthermore, these recent results suggest that sex can be maintained even when very little [[genetic variability]] is produced.
 Prior to 2013, when the "[[Teleomorph, anamorph and holomorph#One fungus, one name|one fungus, one name]]" 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==
+== References ==
-{{Reflist|colwidth=30em|refs=
+{{Reflist|colwidth=30em}}
+== Further reading ==
-<ref name="Mrázek2015">{{cite journal |last1=Mrázek |first1=J |last2=Pachlová |first2=V |last3=Buňka |first3=F |last4=Černíková |first4=M |last5=Dráb |first5=V |last6=Bejblová |first6=M |last7=Staněk |first7=K |last8=Buňková |first8=L |title=Effects of different strains Penicillium nalgiovense in the Nalžovy cheese during ripening|journal=Journal of the Science of Food and Agriculture |volume=96 |issue=7 |pages=2547–54 |pmid=26251231|doi=10.1002/jsfa.7375|date=11 September 2015 }}</ref>
+{{refbegin}}
+* {{cite book | vauthors = Harshberger JW |title=A Text-Book of Mycology and Plant Pathology |url=https://archive.org/details/atextbookmycolo00harsgoog |location=London |publisher=Churchill Livinstone |year=1917 |oclc=879097707}}
-<ref name=Baigre2003>{{Cite book |last=Balgrie |first=B. |title=Taints and Off-flavours in Food |year=2003 |publisher=CRC Press |isbn=978-1-85573-449-4 |page=134 |url=https://books.google.com/books?id=xDcNCY31oQ4C&pg=PA134 |access-date=2013-02-03}}</ref>
+* {{cite journal | vauthors = Asan A |year=2004 |title=''Aspergillus'', ''Penicillium'', and Related Species Reported from Turkey |journal=Mycotaxon |volume=89 |issue=1 |pages=155–7 |url=http://www.mycotaxon.com/resources/checklists/asan-v89-checklist.pdf }}
+* {{cite book |url=https://books.google.com/books?id=3KJewsIL5vQC&q=genus+penicillium&pg=PA66 |last1=Samson |first1=R.A. |first2=J.I. |last2=Pitt |title=Integration of Modern Taxonomic Methods For Penicillium and Aspergillus Classification |isbn=978-9058231598 |publisher=CRC Press |year=2000 |page=66 }}
-<ref name=Chang1995>{{cite journal |last1=Chang |first1=JCS |last2=Foarde |first2=KK |last3=Vanosdell |first3=DW. |title=Growth evaluation of fungi (''Penicillium'' and ''Aspergillus spp.'') on ceiling tiles |journal=Atmospheric Environment |year=1995 |volume=29 |issue=17 |pages=2331 37 |doi=10.1016/1352-2310(95)00062-4|bibcode=1995AtmEn..29.2331C }}</ref>
+{{refend}}
-<ref name=daCosta1998>{{cite journal |last1=da Costa |first1=GL |last2=de Moraes |first2=AM |last3=de Oliveira |first3=PC |title=Pathogenic action of ''Penicillium'' species on mosquito vectors of human tropical diseases |journal=Journal of Basic Microbiology |year=1998 |volume=38 |issue=5–6 |pages=337–41 |pmid=9871331 |doi=10.1002/(SICI)1521-4028(199811)38:5/6<337::AID-JOBM337>3.0.CO;2-N|s2cid=221867835 }}</ref>
-<ref name=DeCarli1988>{{cite journal |last1=De Carli |first1=L. |last2=Larizza |first2=L. |title=Griseofulvin |journal=Mutation Research |year=1988 |volume=195 |issue=2 |pages=91–126 |pmid=3277037 |doi=10.1016/0165-1110(88)90020-6}}</ref>
-<ref name=Fairs2010>{{cite journal |title=Guidelines on ambient intramural airborne fungal spores |journal=Journal of Investigational Allergology and Clinical Immunology |year=2010 |volume=20 |issue=6 |pages=490–98 |pmid=21243933 |last1=Fairs |first1=A. |last2=Wardlaw |first2=AJ |last3=Thompson |first3=JR |last4=Pashley |first4=CH}}</ref>
-<ref name=Haubrich2003>{{cite book |last=Haubrich |first=W.S. |title=Medical Meanings: A Glossary of Word Origins |edition=2nd |year=2003 |publisher=American College of Physicians |location=Philadelphia, Pennsylvania |isbn=978-1-930513-49-5 |page=175 |url=https://books.google.com/books?id=NXmlIwkQBLAC&pg=PA175 |access-date=2013-02-03}}</ref>
-<ref name=Kirk2008>{{cite book |last1=Kirk |first1=PM |last2=Cannon |first2=PF |last3=Minter |first3=DW |last4=Stalpers |first4=JA |title=Dictionary of the Fungi |url=https://archive.org/details/dictionaryfungit00kirk |url-access=limited |edition=10th |publisher=CABI |location=Wallingford, UK |year=2008 |page=[https://archive.org/details/dictionaryfungit00kirk/page/n515 505] |isbn=978-0-85199-826-8}}</ref>
-<ref name=Leitao2009>{{cite journal |last=Leitão |first=A.L. |title=Potential of ''Penicillium'' species in the bioremediation field |journal=International Journal of Environmental Research and Public Health |year=2009 |volume=6 |issue=4 |pages=1393–417 |pmid=19440525 |doi=10.3390/ijerph6041393 |pmc=2681198|doi-access=free }}</ref>
-<ref name=Link1809>{{cite journal |last=Link |first=JHF |title=Observationes in ordines plantarum naturales. Dissertatio I |journal=Magazin der Gesellschaft Naturforschenden Freunde Berlin |year=1809 |volume=3 |pages=3–42 |language=la|url=http://gdz.sub.uni-goettingen.de/en/dms/loader/img/?PID=PPN608227714_0003&physid=PHYS_0031}}</ref>
-<ref name=Marianski2009>{{cite book |last1=Marianski |first1=S. |last2=Marianski |first2=A. |title=The Art of Making Fermented Sausages |year=2009 |publisher=Bookmagic |location=Seminole, Florida |isbn=978-0-9824267-1-5 |url=https://books.google.com/books?id=3uzSvCdVqYwC&pg=PA47 |page=47 |access-date=2013-02-03}}</ref>
-<ref name=Nicoletti2008>{{cite journal |last1=Nicoletti |first1=R. |last2=Buommino |first2=E. |last3=De Filippis |first3=A. |last4=Lopez-Gresa |first4=M. |last5=Manzo |first5=E. |last6=Carella |first6=A |last7=Petrazzuolo |first7=M |last8=Tufano |first8=M.A. |title=Bioprospecting for antagonistic ''Penicillium'' strains as a resource of new antitumor compounds |journal=  World Journal of Microbiology and Biotechnology|year=2009 |volume=24 |issue=2 |pages=185–95 |doi= 10.1007/s11274-007-9455-y|s2cid=86101370 }}</ref>
-<ref name=Nicoletti2009>{{cite journal |vauthors=Nicoletti R, Manzo E, Ciavatta ML |title=Occurrence and bioactivities of funicone-related compounds |journal=International Journal of Molecular Sciences |year=2009 |volume=10 |issue=4 |pages=1430–44 |pmid=19468317 |pmc=2680625 |doi=10.3390/ijms10041430|doi-access=free }}</ref>
-<ref name=Pitt1979>{{cite book |last=Pitt |first=J.I. |title=The genus ''Penicillium'' and its teleomorphic states ''Eupenicillium'' and ''Talaromyces'' |year=1979 |publisher=Academic Press |isbn=978-0-12-557750-2}}</ref>
-<ref name=Pitt2000>{{cite journal |vauthors=Pitt JI, Basílico JC, Abarca ML, López C |title=Mycotoxins and toxigenic fungi |journal=Medical Mycology |year=2000 |volume=38 |issue=Suppl 1 |pages=41–46 |pmid=11204163 |doi=10.1080/714030911}}</ref>
-<ref name=Rifkind2005>{{cite book |last1=Rifkind |first1=D. |last2=Freeman |first2=G. |title=The Nobel Prize Winning Discoveries in Infectious Diseases |year=2005 |publisher=Academic Press |location=London, UK |pages=43–46 |url=https://books.google.com/books?id=d3wdy3b9VUkC&pg=PA43 |isbn=978-0-12-369353-2 |access-date=2013-02-03}}</ref>
-<ref name=Samson1985>{{cite book |last1=Samson |first1=R.A. |last2=Pitt |first2=J.I. |title=Advances in ''Penicillium'' and ''Aspergillus'' Systematics |year=1985 |publisher=Springer |isbn=978-0-306-42222-5 |url-access=registration |url=https://archive.org/details/advancesinpenici0000inte }}</ref>
-<ref name=Samson2004>{{cite journal |vauthors=Samson RA, Seifert KA, Kuijpers AF, Houbraken JA, Frisvad JC |title=Phylogenetic analysis of ''Penicillium'' subgenus ''Pencillium'' using partial beta-tubulin sequences |journal=Studies in Mycology |year=2004 |volume=49 |pages=175–200 |url=http://www.cbs.knaw.nl/publications/1049/part5.pdf }}</ref>
-<ref name=Semenov2003>{{cite book |vauthors=Semenov SA, Gumargalieva KZ, Zaikov GE |title=Biodegradation and Durability of Materials Under the Effect of Microorganisms (New Concepts in Polymer Science) |publisher=V.S.P. Intl Science |isbn=978-90-6764-388-7 |pages=34–35 |url=https://books.google.com/books?id=Kj89NRHHqPMC&pg=PA35 |date=2003-08-01}}</ref>
-<ref name=Singh2008>{{cite journal |vauthors=Singh P, Rathinasamy K, Mohan R, Panda D |title=Microtubule assembly dynamics: an attractive target for anticancer drugs |journal=IUBMB Life |year=2008 |volume=60 |issue=6 |pages=368–75 |pmid=18384115 |doi=10.1002/iub.42|s2cid=111334 |doi-access=free }}</ref>
-<ref name="urlMycoBank: Penicillium">{{cite web |title=''Penicillium'' Link 1809 |url=http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=9257 |publisher=International Mycological Association |work=MycoBank |access-date=2011-07-14}}</ref>
-<!-- <ref name=Ustianowski2008>{{cite journal |vauthors=Ustianowski AP, Sieu TP, Day JN |title=''Penicillium marneffei'' infection in HIV |journal=Current Opinion in Infectious Diseases |year=2008 |volume=21 |issue=1 |pages=31–36 |pmid=18192783 |doi=10.1097/QCO.0b013e3282f406ae}}</ref> -->
-<ref name=Valdez2006>{{cite journal |vauthors=Valdez JG, Makuch MA, Ordovini AF, Masuelli RW, Overy DP, Piccolo RJ |title=First report of ''Penicillium allii'' as a field pathogen of garlic (''Allium sativum'') |journal=Plant Pathology |year=2006 |volume=55 |issue=4 |page=583 |doi=10.1111/j.1365-3059.2006.01411.x|doi-access=free }}</ref>}}
-* {{cite book |last=Harshberger |first=J. W. |title=A Text-Book of Mycology and Plant Pathology |url=https://archive.org/details/atextbookmycolo00harsgoog |location=London |publisher=Churchill Livinstone |year=1917 |oclc=879097707}}
 == External links ==
 {{Sister project links|d=Q843136|c=category:Penicillium|n=no|wikt=yes|b=no|v=no|voy=no|s=no|m=no|mw=no|q=no|species=yes}}
-* {{cite journal |last=Asan |first=A. |year=2004 |title=''Aspergillus'', ''Penicillium'', and Related Species Reported from Turkey |journal=Mycotaxon |volume=89 |issue=1 |pages=155–7 |url=http://www.mycotaxon.com/resources/checklists/asan-v89-checklist.pdf }}
-* {{cite book |url=https://books.google.com/books?id=3KJewsIL5vQC&q=genus+penicillium&pg=PA66 |last1=Samson |first1=R.A. |first2=J.I. |last2=Pitt |title=Integration of Modern Taxonomic Methods For Penicillium and Aspergillus Classification |isbn=978-9058231598 |publisher=CRC Press |year=2000 |page=66 }}
 {{Taxonbar|from=Q843136}}