Penicillium: Difference between revisions

| type_species_authority = Link (181809)

| subdivision = [[List of Penicillium species|over 300]]

| synonyms =''Floccaria'' <small>[[Grev.]]{{Collapsible (1827)</small><br/>list|bullets=on

|''AspergilloidesAspergillopsis'' <small>[[R. P. Dierckx|Dierckx]]Sopp (19011912)</small><br/>

|''WalziaCarpenteles'' <small>[[Nikolai Vasilyevich Sorokin|Sorokin]]Langeron (18711922)</small><br/>

|''PritzeliellaChromocleista'' <small>[[Henn.]]Yaguchi & Udagawa (19031993)</small>

'''''Penicillium''''' ({{IPAc-en|ˌ|p|ɛ|n|ɪ|ˈ|s|ɪ|l|i|ə|məm}}) is a [[genus]] of [[Ascomycota|ascomycetous]] [[Fungusfungus|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>{{Cite namebook | 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=Kirk2008en}}</ref>

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>Identification{{cite andjournal nomenclature| ofvauthors the= genusVisagie PenicilliumCM, C.M.Houbraken Visagie1J, J.Frisvad Houbraken1JC, ,Hong SB, J.C.Klaassen Frisvad2CH, ,Perrone G, S.-B.Seifert Hong3KA, C.H.W.Varga Klaassen4J, G.Yaguchi Perrone5T, K.A.Samson Seifert6,RA J.| Varga7,title T.= Yaguchi8,Identification R.A.and Samson,nomenclature 22of Septemberthe genus Penicillium | journal = Studies in Mycology | volume = 78 | issue = 1 | pages = 343–371 | date = June 2014, https://dx.| pmid = 25505353 | pmc = 4261876 | doi.org/ = 10.1016/j.simyco.2014.09.001 }}</ref><ref>{{Cite book | vauthors name=Link1809 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 nameconference |book-title=Samson1985Advances 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>{{Cite namebook | 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=Pitt1979en}}</ref> Species included in subgenus ''Biverticillium'' were later merged into ''[[Talaromyces]]''.

Selected species include;:

* ''[[Penicillium bilaiae]]'', which is an agricultural inoculant

* ''[[Penicillium camemberti]]'', which is used in the production of [[Camembert (cheese)|Camembert]] and, [[Brie cheese|Brie]] and [[cheeseCambozola]]s cheeses

* ''[[Penicillium chrysogenum]]'' (previously known as ''[[Penicillium notatum]]''), which produces the [[antibiotic]] [[penicillin]]

* ''[[Penicillium expansum]]'', a pathogen of apples and other fruit, produces [[patulin]]

* ''[[Penicillium roqueforti]]'', which is used in making [[Roquefort cheese|Roquefort]], [[Danish Blue cheese]], English Blue [[Stilton cheese]], and [[Gorgonzola cheese]], and [[Cambozola]]

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 namebook | 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=Haubrich2003en}}</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 |last vauthors = Pitt|first= J. |date=1985 |title=A laboratory guide to common Penicillium species |urljournal=https://www.semanticscholar.org/paper/A-laboratory-guide-to-common-Penicillium-species-Pitt/7c86b2f5283c16d926fa93de64ef535040eec860?p2dfMycologia | volume=79 |page=491 |doi=10.2307/3807483 |jstor=3807483 |s2cid=84610634}}</ref>

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 name=Samson2004 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 name=Pitt2000 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>{{Cite book | vauthors name=Baigre2003 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 name=Valdez2006 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 name=daCosta1998 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>{{cite journal | vauthors name=Fairs2010 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 name=Chang1995 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>{{Cite namebook | 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=Semenov2003en}}</ref>

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 [[Blueblue 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 name="Mrázek2015" 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 name=Marianski2009 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>{{cite journal | vauthors name=Leitao2009 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 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 namebook | 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=Rifkind2005en}}</ref>

[[Griseofulvin]] is an [[antifungal drug]] and a potential [[chemotherapeutic agent]]<ref>{{cite journal | vauthors name=Singh2008 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 name=DeCarli1988 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 namejournal |vauthors=Nicoletti2009Nicoletti 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=Nicoletti2008":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=Nicoletti2008":0" />

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 |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.Proceedings Acad.of Sci.the U.S.A.National Academy of Sciences of the United States of America | volume = 110 | issue = 4 | pages =1476–81 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 date =2008 August 2007 | pmid = 18663385 | pmc = 2488364 | doi = 10.1371/journal.pone.0002879 | doi-access = free | 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.

== References ==

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