|
|
この記事は検証可能な参考文献や出典が全く示されていないか、不十分です。
出典を追加して記事の信頼性向上にご協力ください。（2010年11月） |
| アオカビ |
|
アオカビ（顕微鏡写真、x200）
|
| 分類 |
| 界 |
: |
菌界 Fungi |
| 門 |
: |
子嚢菌門 Ascomycota |
| 綱 |
: |
ユーロチウム菌綱 Eurotiomycetes |
| 目 |
: |
ユーロチウム目 Eurotiales |
| 科 |
: |
マユハキタケ科 Trichocomaceae |
| 属 |
: |
アオカビ属 Penicillium |
|
| 学名 |
| Penicillium Link, 1809 |
| タイプ種 |
Penicillium expansum
Link (1809) |
| 種 |
|
300種以上
|
アオカビやコウジカビを含む様々な無菌培養された菌類
アオカビ（Penicillium）はアオカビ属（ペニシリウム属）に属するカビの総称で、もっとも普遍的に見られる不完全菌の一つである。胞子の色が肉眼で青みを帯びた水色であることからその名がある。ただし、白や緑がかった色のものも見られ、必ずしもすべてのアオカビ属（Penicillium属）のカビが青いわけではない。
顕微鏡で観察すると、筆状体（ひつじょうたい、penicillus）と呼ばれる筆のような形の構造が見られ、その先端に胞子がついているのが判る。これがこのカビの学名の由来になっている。世界で初めての抗生物質であるペニシリンが、この種のカビから発見されたことは有名である。また、ゴルゴンゾーラ、ロックフォールなどの代表的なチーズの製造に用いられるカビもアオカビの仲間である。
目次
- 1 生物学的特徴
- 1.1 系統分類学的位置
- 1.2 アオカビの胞子（分生子）
- 1.3 類似のカビ
- 2 アオカビとヒトの病気
- 3 アオカビによる植物の病気
- 4 アオカビの利用
- 4.1 アオカビとペニシリン
- 4.2 アオカビとチーズ
生物学的特徴
アオカビは最も普遍的に見られるカビの一つであり、常に空中に胞子が飛散しているので、パンや餅などには、真っ先に生えて来る。胞子は発芽すると基質中に菌糸をのばし、コロニーを形成する。すぐに胞子形成を始めるので、コロニーは青みがかった色になる。
系統分類学的位置
カビの系統分類は有性生殖を行うときの世代（完全世代）を基準に行われるが、中には完全世代の状態が見つかっていない（あるいは元から有性生殖を行わないという可能性もある）ものが存在する。これらは不完全菌という（やや便宜的な）分類群として扱われ、その無性生殖世代（不完全世代）の形態的特徴からさらにいくつかに分類されている。不完全菌のうち筆状体を作るものがPenicillium属に分類される。このため、ひとたびPenicillium属として記載された菌であっても完全世代が発見されると、それに見合った分類群に振り分けられ、新たに学名がつけなおされる。
完全世代が確認されたPenicillium属のほとんどは、閉子嚢殻という小さな球形の子実体を形成するが、中にはきのことして扱われる大型の子実体を形成するものもある。これらは子嚢菌のユーロチウム目に分類されているが、Eupenicillium属、Hamigera属、Penicilliopsis属、Talaromyces属、Trichocoma属と幾つもの属にまたがっている。「きのこ」を作るものにはPenicilliopsis属のカキノミタケやTrichocoma属のマユハキタケが知られる。
有性生殖が知られていないものについても、リボソームRNAの相同性から子嚢菌の系統に属することが確認されているものが多い。
アオカビの胞子（分生子）
アオカビなど、不完全菌の胞子は分生子と呼ばれる。アオカビの分生子柄は、基質の表面に短く立ち、先端で幾つかの短い枝に分かれる。それぞれの枝の先にフィアライドと呼ばれる分生子形成細胞が数個並ぶ。フィアライドは紡錘形で、その先端から分生子を出芽するように形成する。分生子が完成すると、分生子とフィアライドの間は切り放され、前の分生子を押し出すようにして新しい分生子が出芽し、次第に分生子の数珠が作られる。
分生子柄の枝は、互いに小さな角度を作り、寄り添うように伸び、それぞれの先端のフィアライドも枝の先端方向を向くので、それぞれのフィアライドの先端の分生子の数珠も同じ方向へ伸び、全体としては筆や箒、松明のような形になる。分生子はどんどん作られ、数珠は長く伸び、やがて崩れるので、コロニー中心部では菌糸体は崩れた胞子の下に埋まってしまう。
フィアライドから分生子の数珠を作る点では、アオカビはコウジカビとよく似ており、完全世代でも類縁関係が確認されている。ちなみに、コウジカビや、その他の不完全菌にも青っぽい胞子を作るものがあるので、色だけで判断するのは無理である。
類似のカビ
肉眼的にはコウジカビが似ている。それ以外のものは、食品などに発生することが多くない。形態的に似ているのはPaecilomycesである。アオカビに似た分生子柄の分枝を持ち、先端のフィアライドから胞子の数珠を出すのも同じであるが、胞子は無色で、コロニーは白く見える。また、Gliocladiumも同じような分生子柄を持つが、これは胞子を鎖ではなく粘液球の形でつける。
アオカビとヒトの病気
ほとんどのアオカビは、健康なヒトには感染せず非病原性である。ただしアオカビの仲間のうち、二形性（菌糸型と酵母様細胞の両方の形態をとる性質）を示すものは比較的毒性が強く、後天性免疫不全症候群(AIDS)患者に日和見感染を起こす例(P. marneffeiによる)が報告されている。また、爪、耳、肺、尿路においてペニシリウム症と呼ばれる感染症を起こすことがある。
アオカビの大部分はカビ毒（マイコトキシン）を産生しないため、これらが直接に重篤な食中毒の原因になることはほとんどない。ただしアオカビが生えた食品では、他の有害なカビの増殖も進んでいると考えるべきである。
アオカビによる植物の病気
アオカビによる植物の病気は「青かび病」という名が付いたものが多い。身近で重要なものでは、P. expansumによるミカン青かび病がある。同じくP. expansumによるものには、リンゴ青かび病、サツマイモ青かび病などがある。
イネの病変米にもアオカビによるものが多数あり、病原菌としてP. cyclopium, P. islandicum, P. commune, P. citro-viride, P. rugulosum, P. citrinum, P. italicumなどが報告されている。黄変米は、これらの菌が収穫後の米の運送時に感染したために生じたものである。
アオカビの利用
アオカビとペニシリン
医学的にアオカビは、病原菌としてよりむしろペニシリンの発見につながったことから重要視されている。ペニシリンはアレクサンダー・フレミングが1928年に発見した世界初の抗生物質である。一説には、他の細菌を培養するための培地にたまたまアオカビ(当時のP. notatum、現在はP. chrysogenum)が混入した際、そのアオカビのコロニー周辺に細菌が生育しない領域（阻止円）が出来ることを見つけたことがその発見につながったと言われる。アオカビにとってペニシリンは、他の生物（特に細菌）との生存競争を有利に進めるために産生するアロモンであると考えられている。その全合成方法が開発されるまでの間、ペニシリンを製造するためアオカビの大量培養が行われた。ペニシリン以外の抗生物質を産生する菌として、グリセオフルビンを産生するP. griseofluvumが見つかっている。
アオカビとチーズ
アオカビのうちある種のものはチーズの製造に用いられる。特にアオカビを内部に植えつけたものはブルーチーズと呼ばれ、ゴルゴンゾーラ(P. galaucumを使用)やロックフォール(P.roquefortiを使用)がその代表として知られる。なおカマンベールチーズやブリーチーズなどは、いわゆる「白カビ」を表面に生やすが、これらも生物学的にはPenicillium属に属するカビである。これらのチーズ製造過程で、アオカビはチーズを発酵・熟成させるとともに、独特の風味・香りをチーズに与える。
| Penicillium |
|
| Penicillium sp. |
| Scientific classification |
| Kingdom: |
Fungi |
| Phylum: |
Ascomycota |
| Class: |
Eurotiomycetes |
| Order: |
Eurotiales |
| Family: |
Trichocomaceae |
| Genus: |
Penicillium
Link (1809) |
| Type species |
Penicillium expansum
Link (1809) |
| Species |
|
over 300
List of Penicillium species
|
| Synonyms[1] |
|
Floccaria Grev. (1827)
Aspergilloides Dierckx (1901)
Walzia Sorokin (1871)
Pritzeliella Henn. (1903)
|
Penicillium () is a genus of ascomycetous fungi of major importance in the natural environment as well as 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 inside the body. Other species are used in cheesemaking. According to the Dictionary of the Fungi (10th edition, 2008), the widespread genus contains over 300 species.[2]
Contents
- 1 Taxonomy
- 1.1 Species
- 1.2 Etymology
- 2 Characteristics
- 3 Ecology
- 4 Economic value
- 5 Reproduction
- 6 References
- 7 External links
Taxonomy
The genus was first described in the scientific literature by Johann Heinrich Friedrich Link in his 1809 work Observationes in ordines plantarum naturales.[3] Link included three species—P. candidum, P. expansum, and P. glaucum—all of which produced a brush-like conidiophore (asexual fruiting structure). The common apple rot fungus P. expansum was selected as the type species.[4]
In a 1979 monograph, John I. Pitt divided Penicillium into four subgenera based on conidiophore morphology and branching pattern: Aspergilloides, Biverticillium, Furcatum, and Penicillium.[5]
Penicillium is classified as a genus of: domain eukaryota, kingdom Fungi, division Ascomycota (order Eurotiales, class Eurotiomycetes, family Trichocomaceae).[6]
Species
Main article: List of Penicillium species
Various fungi including
Penicillium and
Aspergillus species growing in axenic culture
Some penicillium mold on mandarin oranges
Selected species include;
- Penicillium albocoremium
- Penicillium aurantiogriseum
- Penicillium bilaiae, which is an agricultural inoculant
- Penicillium camemberti, which is used in the production of Camembert and Brie 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 plant pathogen
- Penicillium echinulatum produces Mycophenolic acid
- Penicillium expansum, a plant pathogen
- Penicillium funiculosum, a plant pathogen
- Penicillium glabrum
- Penicillium glaucum, which is used in making Gorgonzola cheese
- Penicillium italicum
- Penicillium lacussarmientei
- Penicillium marneffei, a thermally dimorphic species endemic in Southeast Asia, which presents a threat of systemic infection to AIDS patients
- Penicillium purpurogenum
- Penicillium roqueforti, which is used in making Roquefort, Danish Blue cheese, and also recently Gorgonzola
- Penicillium stoloniferum
- Penicillium ulaiense
- Penicillium verrucosum produces ochratoxin A
- Penicillium viridicatum produces ochratoxin
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
Penicillium Spp. under bright field microscopy (10 x 100 magnification) with lactophenol cotton blue stain
The thallus (mycelium) typically consists of a highly branched network of multinucleate, septate, usually colorless hyphae. Many-branched conidiophores sprout on the mycelia, bearing individually constricted conidiospores. The conidiospores are the main dispersal route of the fungi, and often are green in color.
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 as molds, they are among the main causes of food spoilage, especially species of subgenus Penicillium.[8] 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.[9] 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;[10] and P. allii, garlic.[11] 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.[12] P. marneffei, which causes mortality in the Vietnamese bamboo rats, has become a common opportunistic infection of HIV-infected individuals in southeast Asia.[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, P. steckii, P. notatum, 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
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 to improve the taste of sausages and hams, and to prevent colonization by other molds and bacteria.[17]
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 because of their ability to break down a variety of xenobiotic compounds.[18]
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 Boris 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.[19]
Griseofulvin is an antifungal drug and a potential chemotherapeutic agent[20] that was discovered in P. griseofulvum.[21] Additional species that produce compounds capable of inhibiting the growth of tumor cells in vitro include: P. pinophilum,[22] P. canescens,[23] and P. glabrum.[23]
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.[24] 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.[25] finally demonstrated sexual reproduction in P. chrysogenum.
Penicillium marneffei, an AIDS-associated pathogen, was also previously assumed to reproduce exclusively by asexual means. This assumption was largely based on the highly clonal population structure of this species. However, recent work has revealed that the genes required for meiosis are present in P. marneffei.[26] This and other evidence indicated that mating and genetic recombination does occur in this species. It was concluded that P. marneffei is sexually reproducing, but recombination is most likely to occur across spatially and genetically limited distances in natural populations resulting in a highly clonal population structure.[26]
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][28] Furthermore, these recent results suggest that sex can be maintained even when very little genetic variability is produced.
References
- ^ "Penicillium Link 1809". MycoBank. International Mycological Association. Retrieved 2011-07-14.
- ^ Kirk, PM; Cannon, PF; Minter, DW; Stalpers, JA (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CABI. p. 505. ISBN 978-0-85199-826-8.
- ^ Link, JHF (1809). "Observationes in ordines plantarum naturales. Dissertatio I". Magazin der Gesellschaft Naturforschenden Freunde Berlin (in Latin) 3: 3–42.
- ^ Samson, R.A.; Pitt, J.I. (1985). Advances in Penicillium and Aspergillus Systematics. Springer. ISBN 978-0-306-42222-5.
- ^ Pitt, J.I. (1979). The genus Penicillium and its teleomorphic states Eupenicillium and Talaromyces. Academic Press. ISBN 978-0-12-557750-2.
- ^ "Names record: Penicillium Link". Index Fungorum. CAB International. Retrieved 2011-07-14.
- ^ Haubrich, W.S. (2003). Medical Meanings: A Glossary of Word Origins (2nd ed.). Philadelphia, Pennsylvania: American College of Physicians. p. 175. ISBN 978-1-930513-49-5. Retrieved 2013-02-03.
- ^ Samson RA, Seifert KA, Kuijpers AF, Houbraken JA, Frisvad JC. (2004). "Phylogenetic analysis of Penicillium subgenus Pencillium using partial beta-tubulin sequences" (PDF). Studies in Mycology 49: 175–200.
- ^ Pitt JI, Basílico JC, Abarca ML, López C.; Basílico; Abarca; López (2000). "Mycotoxins and toxigenic fungi". Medical Mycology 38 (Suppl 1): 41–46. doi:10.1080/714030911. PMID 11204163.
- ^ Balgrie, B. (2003). "Taints and Off-flavours in Food". CRC Press. p. 134. ISBN 978-1-85573-449-4. Retrieved 2013-02-03.
- ^ Valdez JG, Makuch MA, Ordovini AF, Masuelli RW, Overy DP, Piccolo RJ.; Makuch; Ordovini; Masuelli; Overy; Piccolo (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.
- ^ 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): 337–41. doi:10.1002/(SICI)1521-4028(199811)38:5/6<337::AID-JOBM337>3.0.CO;2-N. PMID 9871331.
- ^ Ustianowski AP, Sieu TP, Day JN.; Sieu; Day (2008). "Penicillium marneffei infection in HIV". Current Opinion in Infectious Diseases 21 (1): 31–36. doi:10.1097/QCO.0b013e3282f406ae. PMID 18192783.
- ^ Fairs, A.; Wardlaw, AJ; Pashley, CH (2010). "Guidelines on ambient intramural airborne fungal spores". Journal of Investigational Allergology and Clinical Immunology 20 (6): 490–98. PMID 21243933.
- ^ Chang, JCS; Foarde, KK; Vanosdell, DW. (1995). "Growth evaluation of fungi (Penicillium and Aspergillus spp.) on ceiling tiles". Atmospheric Environment 29 (17): 2331 37. doi:10.1016/1352-2310(95)00062-4.
- ^ Semenov SA, Gumargalieva KZ, Zaikov GE. (2003-08-01). Biodegradation and Durability of Materials Under the Effect of Microorganisms (New Concepts in Polymer Science). V.S.P. Intl Science. pp. 34–35. ISBN 978-90-6764-388-7.
- ^ Marianski, S.; Marianski, A. (2009). The Art of Making Fermented Sausages. Seminole, Florida: Bookmagic. p. 47. ISBN 978-0-9824267-1-5. Retrieved 2013-02-03.
- ^ Leitão, A.L. (2009). "Potential of Penicillium species in the bioremediation field" (PDF). International Journal of Environmental Research and Public Health 6 (4): 1393–417. doi:10.3390/ijerph6041393. PMC 2681198. PMID 19440525.
- ^ Rifkind, D.; Freeman, G. (2005). The Nobel Prize Winning Discoveries in Infectious Diseases. London, UK: Academic Press. pp. 43–46. ISBN 978-0-12-369353-2. Retrieved 2013-02-03.
- ^ Singh P, Rathinasamy K, Mohan R, Panda D.; Rathinasamy; Mohan; Panda (2008). "Microtubule assembly dynamics: an attractive target for anticancer drugs". IUBMB Life 60 (6): 368–75. doi:10.1002/iub.42. PMID 18384115.
- ^ De Carli, L.; Larizza, L. (1988). "Griseofulvin". Mutation Research 195 (2): 91–126. doi:10.1016/0165-1110(88)90020-6. PMID 3277037.
- ^ Nicoletti R, Manzo E, Ciavatta ML.; Manzo; Ciavatta (2009). "Occurrence and bioactivities of funicone-related compounds". International Journal of Molecular Sciences 10 (4): 1430–44. doi:10.3390/ijms10041430. PMC 2680625. PMID 19468317.
- ^ a b Nicoletti, R.; Buommino, E.; De Filippis, A.; Lopez-Gresa, M.; Manzo, E.; Carella, A; Petrazzuolo, M; Tufano, M.A. (2009). "Bioprospecting for antagonistic Penicillium strains as a resource of new antitumor compounds". World Journal of Microbiology 24 (2): 185–95. doi:10.1007/s11274-007-9455-y.
- ^ Ropars J, Dupont J, Fontanillas E, Rodríguez de la Vega RC, Malagnac F, Coton M, Giraud T, López-Villavicencio M; Dupont; Fontanillas; Rodríguez de la Vega RC; Malagnac; Coton; Giraud; López-Villavicencio (2012). "Sex in cheese: evidence for sexuality in the fungus Penicillium roqueforti". PLoS ONE 7 (11): e49665. doi:10.1371/journal.pone.00496. PMC 3504111. PMID 23185400.
- ^ Böhm J, Hoff B, O'Gorman CM, Wolfers S, Klix V, Binger D, Zadra I, Kürnsteiner H, Pöggeler S, Dyer PS, Kück U; Hoff; O'Gorman; Wolfers; Klix; Binger; Zadra; Kürnsteiner; Pöggeler; Dyer; Kück (January 2013). "Sexual reproduction and mating-type-mediated strain development in the penicillin-producing fungus Penicillium chrysogenum". Proc. Natl. Acad. Sci. U.S.A. 110 (4): 1476–81. doi:10.1073/pnas.1217943110. PMC 3557024. PMID 23307807.
- ^ a b Henk DA, Shahar-Golan R, Devi KR, Boyce KJ, Zhan N, Fedorova ND, Nierman WC, Hsueh PR, Yuen KY, Sieu TP, Kinh NV, Wertheim H, Baker SG, Day JN, Vanittanakom N, Bignell EM, Andrianopoulos A, Fisher MC; Shahar-Golan; Devi; Boyce; Zhan; Fedorova; Nierman; Hsueh; Yuen; Sieu; Kinh; Wertheim; Baker; Day; Vanittanakom; Bignell; Andrianopoulos; Fisher (2012). "Clonality despite sex: the evolution of host-associated sexual neighborhoods in the pathogenic fungus Penicillium marneffei". PLoS Pathog. 8 (10): e1002851. doi:10.1371/journal.ppat.1002851. PMC 3464222. PMID 23055919.
- ^ Malik SB, Pightling AW, Stefaniak LM, Schurko AM, Logsdon JM; Pightling; Stefaniak; Schurko; Logsdon Jr (2008). "An expanded inventory of conserved meiotic genes provides evidence for sex in Trichomonas vaginalis". PLoS ONE 3 (8): e2879. doi:10.1371/journal.pone.0002879. PMC 2488364. PMID 18663385.
- ^ Bernstein H and Bernstein C (2013). Evolutionary Origin and Adaptive Function of Meiosis. In Meiosis: Bernstein C and Bernstein H, editors. ISBN 978-953-51-1197-9, InTech, http://www.intechopen.com/books/meiosis/evolutionary-origin-and-adaptive-function-of-meiosis
- Harshberger, J.W. (1917). A Text-Book of Mycology and Plant Pathology. Churchill Livinstone.
External links
|
Wikimedia Commons has media related to Penicillium. |
|
Wikisource has the text of the 1921 Collier's Encyclopedia article Penicillium. |
- Samson, R. A. & J I Pitt Integration of Modern Taxonomic Methods For Penicillium and Aspergillus Classification (with an illustration on the cover)
- Asan, A. (2004, Last updated: February, 2015). Aspergillus, Penicillium, and Related Species Reported from Turkey. Mycotaxon. 89 (1): 155-157. Link: http://www.mycotaxon.com/resources/checklists/asan-v89-checklist.pdf
