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Agaricomycetes

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The Agaricomycetes are a class of fungi in the division Basidiomycota. The taxon is roughly identical to that defined for the Homobasidiomycetes (alternatively called holobasidiomycetes) by Hibbett & Thorn,[2] with the inclusion of Auriculariales and Sebacinales. It includes not only mushroom-forming fungi, but also most species placed in the deprecated taxa Gasteromycetes and Homobasidiomycetes.[3] Within the subdivision Agaricomycotina, which already excludes the smut and rust fungi, the Agaricomycetes can be further defined by the exclusion of the classes Tremellomycetes and Dacrymycetes, which are generally considered to be jelly fungi. However, a few former "jelly fungi", such as Auricularia, are classified in the Agaricomycetes. According to a 2008 estimate, Agaricomycetes include 17 orders, 100 families, 1147 genera, and about 21000 species.[4] Modern molecular phylogenetic analyses have been since used to help define several new orders in the Agaricomycetes: Amylocorticiales, Jaapiales,[5] Stereopsidales,[6] and Lepidostromatales.[7]

Classification

Although morphology of the mushroom or fruit body (basidiocarp) was the basis of early classification of the Agaricomycetes,[8] this is no longer the case. As an example, the distinction between the Gasteromycetes (including puffballs) and Agaricomycetes (most other agaric mushrooms) is no longer recognized as a natural one—various puffball species have apparently evolved independently from agaricomycete fungi. However, most mushroom guide books still group the puffballs or gasteroid forms separate from other mushrooms because the older Friesian classification is still convenient for categorizing fruit body forms. Similarly, modern classifications divide the gasteroid order Lycoperdales between Agaricales and Phallales.

Features

All members of the class produce basidiocarps which range in size from tiny cups a few millimeters across to a giant polypore (Phellinus ellipsoideus) greater than several meters across and weigh up to 500 kilograms (1,100 lb).[9] The group also includes what are arguably the largest and oldest individual organisms on earth: the mycelium of one individual Armillaria gallica has been estimated to extend over 150,000 square metres (37 acres) with a mass of 10,000 kg (22,000 lb) and an age of 1,500 years.[10]

Ecology

Nearly all species are terrestrial (a few are aquatic), occurring in a wide range of environments where most function as decayers, especially of wood. However, some species are pathogenic or parasitic, and yet others are symbiotic (i.e., mutualistic), these including the important ectomycorrhizal symbionts of forest trees. General discussions on the forms and life cycles of these fungi are developed in the article on mushrooms, in the treatments of the various orders (links in table at right), and in individual species accounts.

Evolution

A study of 5,284 species with a backbone phylogeny based on 104 genomes[11] has suggested the following dates of evolution:

Agaricomycetidae ~185 million years ago (174 million years ago192 million years ago)
Cantharellales 184 million years ago  (144 million years ago261 million years ago)
Agaricales 173 million years ago  (160 million years ago-182 million years ago)
Hymenochaetales 167 million years ago (130 million years ago180 million years ago)
Boletales 142 million years ago (133 million years ago153 million years ago)

Fossil record

The fruit bodies of Agaricomycetes are extremely rare in the fossil record, and the class does not yet pre-date the Early Cretaceous (146–100 Ma).[12] The oldest Agaricomycetes fossil, dating from the lower Cretaceous (130–125 Ma) is Quatsinoporites. It is a fragment of a poroid fruit body with features that suggest it could be a member of the family Hymenochaetaceae.[13] Based on molecular clock analysis, the Agaricomycetes are estimated to be about 290 million years old.[14]

Phylogeny

Modern molecular phylogenetics suggest the following relationships:[15]

   

Basidiomycetes (outgroup)

  Agaricomycetes  

Cantharellales

     

Sebacinales

     

Auriculariales

       

Stereopsidales

Phallomycetidae

Geastrales

     

Hysterangiales

     

Gomphales

   

Phallales

             

Trechisporales

     

Hymenochaetales

       

Thelephorales

   

Polyporales

         

Corticiales

     

Jaapiales

   

Gloeophyllales

         

Russulales

Agaricomycetidae

Agaricales

     

Boletales

     

Amylocorticiales

     

Lepidostromatales

   

Atheliales

                             

Genera incertae sedis

There are many genera in the Agaricomycetes that have not been classified in any order or family. These include:

References

  1. ^ Doweld A. (2001). Prosyllabus Tracheophytorum, Tentamen systematis plantarum vascularium (Tracheophyta) [An attempted system of the vascular plants]. Moscow, Russia: GEOS. pp. 1–111. ISBN 978-5-89118-283-7.
  2. ^ Hibbett DS, Thorn RG (2001). McLaughlin DJ, et al. (eds.). The Mycota, Vol. VII. Part B., Systematics and Evolution. Berlin, Germany: Springer-Verlag. pp. 121–168.
  3. ^ Hibbett DS; et al. (2007). "A higher level phylogenetic classification of the Fungi". Mycological Research. 111 (5): 509–547. CiteSeerX 10.1.1.626.9582. doi:10.1016/j.mycres.2007.03.004. PMID 17572334.
  4. ^ Kirk PM, Cannon PF, Minter DW, Stalpers JA (2008). Dictionary of the Fungi (10th ed.). Wallingford, UK: CAB International. pp. 12–13. ISBN 978-0-85199-826-8.
  5. ^ Binder M, Larsson KH, Matheny PB, Hibbett DS (2010). "Amylocorticiales ord. nov. and Jaapiales ord. nov.: Early diverging clades of Agaricomycetidae dominated by corticioid forms". Mycologia. 102 (4): 865–880. doi:10.3852/09-288. PMID 20648753.
  6. ^ Sjökvist E, Pfeil BE, Larsson E, Larsson K-H (2014). "Stereopsidales – a new order of mushroom-forming fungi". PLOS ONE. 9 (8): e106204. Bibcode:2014PLoSO...995227S. doi:10.1371/journal.pone.0095227. PMC 4002437. PMID 24777067. open access
  7. ^ Hodkinson BP, Moncada B, Lücking R (2014). "Lepidostromatales, a new order of lichenized fungi (Basidiomycota, Agaricomycetes), with two new genera, Ertzia and Sulzbacheromyces, and one new species, Lepidostroma winklerianum". Fungal Diversity. 64 (1): 165–179. doi:10.1007/s13225-013-0267-0.
  8. ^ Fries EM (1874). Hymenomycetes Europaei (in Latin). Uppsala: Typis Descripsit Ed. Berling. p. 1.
  9. ^ Cui B-K, Dai Y-C (2011). "Fomitiporia ellipsoidea has the largest fruiting body among the fungi". Fungal Biology. 115 (9): 813–814. doi:10.1016/j.funbio.2011.06.008. PMID 21872178.
  10. ^ Smith M, Bruhn JH, Anderson JB (1992). "The fungus Armillaria bulbosa is among the largest and oldest living organisms". Nature. 356 (6368): 428–431. Bibcode:1992Natur.356..428S. doi:10.1038/356428a0.
  11. ^ Varga T, Krizsán K, Földi C, Dima B, Sánchez-García M, Sánchez-Ramírez S, Szöllősi GJ, Szarkándi JG, Papp V, Albert L, Andreopoulos W, Angelini C, Antonín V, Barry KW, Bougher NL, Buchanan P, Buyck B, Bense V, Catcheside P, Chovatia M, Cooper J, Dämon W, Desjardin D, Finy P, Geml J, Haridas S, Hughes K, Justo A, Karasiński D, Kautmanova I, Kiss B, Kocsubé S6, Kotiranta H, LaButti KM, Lechner BE, Liimatainen K, Lipzen A, Lukács Z, Mihaltcheva S, Morgado LN, Niskanen T, Noordeloos ME, Ohm RA, Ortiz-Santana B, Ovrebo C, Rácz N, Riley R, Savchenko A, Shiryaev A, Soop K, Spirin V, Szebenyi C, Tomšovský M, Tulloss RE, Uehling J, Grigoriev IV, Vágvölgyi C, Papp T, Martin FM, Miettinen O, Hibbett DS, Nagy LG (2019) Megaphylogeny resolves global patterns of mushroom evolution. Nat Ecol Evol
  12. ^ Kiecksee, Anna Philie; Seyfullah, Leyla J.; Dörfelt, Heinrich; Heinrichs, Jochen; Süß, Herbert; Schmidt, Alexander R. (2012). "Pre-Cretaceous Agaricomycetes yet to be discovered: Reinvestigation of a putative Triassic bracket fungus from southern Germany". Fossil Record. 15 (2): 85–89. doi:10.1002/mmng.201200006.
  13. ^ Smith, S.Y.; Currah, R.S.; Stockey, R.A. (2004). "Cretaceous and Eocene poroid hymenophores from Vancouver Island, British Columbia". Mycologia. 96 (1): 180–186. doi:10.2307/3762001. JSTOR 3762001. PMID 21148842.
  14. ^ Floudas D.; Binder, M.; Riley, R.; Barry, K.; Blanchette, R.A.; Henrissat, B.; Martínez, AT.; Otillar, R.; Spatafora, J.W.; Yadav, J.S.; Aerts, A.; Benoit, I.; Boyd, A.; Carlson A.; Copeland, A.; Coutinho, P.M.; de Vries, R.P.; Ferreira, P.; Findley, K.; Foster, B.; Gaskell, J.; Glotzer, D.; Górecki, P.; Heitman, J.; Hesse, C.; Hori, C.; Igarashi, K.; Jurgens, J.A.; Kallen, N.; Kersten, P.; Kohler, A.; Kües, U.; Kumar, TK.; Kuo, A.; LaButti, K.; Larrondo, L.F.; Lindquist, E.; Ling, A.; Lombard, V.; Lucas, S.; Lundell, T.; Martin, R.; McLaughlin, D.J.; Morgenstern, I.; Morin, E.; Murat, C.; Nagy, L.G.; Nolan, M.; Ohm, R.A.; Patyshakuliyeva, A.; Rokas, A.; Ruiz-Dueñas, F.J.; Sabat, G.; Salamov, A.; Samejima, M.; Schmutz, J.; Slot, J.C.; St John, F.; Stenlid, J.; Sun, H.; Sun S.; Syed K.; Tsang, A.; Wiebenga A.; Young, D.; Pisabarro, A.; Eastwood, DC.; Martin, F.; Cullen, D.; Grigoriev I.V.; Hibbett, D.S. (2012). "The Paleozoic origin of enzymatic lignin decomposition reconstructed from 31 fungal genomes". Science. 336 (6089): 1715–1719. Bibcode:2012Sci...336.1715F. doi:10.1126/science.1221748. hdl:10261/60626. PMID 22745431.
  15. ^ Hibbett D, Bauer R, Binder M, Giachini AJ, Hosaka K, Justo A, Larsson E, Larsson K-H, Lawrey JD, Miettinen O, Nagy LG, Nilsson RH, Weiss M, Thorn RG (2014). "Agaricomycetes". In McLaughlin DJ, Spatafora JW (eds.). Systematics and Evolution. The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research. 7A (2nd ed.). Berlin, Heidelberg: Springer-Verlag. pp. 373–429. doi:10.1007/978-3-642-55318-9_14. ISBN 978-3-642-55317-2.
  16. ^ Berniccia, Annarosa; Gorjón, Sergio P.; Nakasone, Karen K. (2011). "Arrasia rostrata (Basidiomycota), a new corticioid genus and species from Italy" (PDF). Mycotaxon. 118: 257–264. doi:10.5248/118.257.
  17. ^ Tzean, S.S.; Estey, R.H. (1991). "Geotrichopsis mycoparasitica gen. et sp. nov. (Hyphomycetes), a new mycoparasite". Mycological Research. 95 (12): 1350–1354. doi:10.1016/S0953-7562(09)80383-3.
  18. ^ Hjortstam, Kurt; Ryvarden, Leif (2001). "Corticioid species (Basidiomycotina, Aphyllophorales) from Colombia III". Mycotaxon. 79: 189–200.
  19. ^ Wu, Sheng-Hua; Wang, Dong-Me; Chen, Yu-Ping (2018). "Purpureocorticium microsporum (Basidiomycota) gen. et sp. nov. from East Asia". Mycological Progress. 17 (3): 357–364. doi:10.1007/s11557-017-1362-5.
  20. ^ Chang, TunTschu; Chou, Wen Neng (2003). "Taiwanoporia, a new aphyllophoralean genus". Mycologia. 95 (6): 1215–1218. doi:10.1080/15572536.2004.11833029. JSTOR 3761921. PMID 21149022.

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Agaricomycetes: Brief Summary

provided by wikipedia EN

The Agaricomycetes are a class of fungi in the division Basidiomycota. The taxon is roughly identical to that defined for the Homobasidiomycetes (alternatively called holobasidiomycetes) by Hibbett & Thorn, with the inclusion of Auriculariales and Sebacinales. It includes not only mushroom-forming fungi, but also most species placed in the deprecated taxa Gasteromycetes and Homobasidiomycetes. Within the subdivision Agaricomycotina, which already excludes the smut and rust fungi, the Agaricomycetes can be further defined by the exclusion of the classes Tremellomycetes and Dacrymycetes, which are generally considered to be jelly fungi. However, a few former "jelly fungi", such as Auricularia, are classified in the Agaricomycetes. According to a 2008 estimate, Agaricomycetes include 17 orders, 100 families, 1147 genera, and about 21000 species. Modern molecular phylogenetic analyses have been since used to help define several new orders in the Agaricomycetes: Amylocorticiales, Jaapiales, Stereopsidales, and Lepidostromatales.

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BY1

provided by wikipedia EN

BY1 is a taxonomically unidentified basidiomycete fungus. ITS sequencing has placed it in the Russulales and is referred to as a stereaceous basidiomycete. Chemotaxonomically supporting its placement in this group, it produces fomannoxins and vibralactones. The fungus' mycelia were isolated from dead aspen in Minnesota, USA. It is presumed to decompose wood by white rot.[1]

The mycelium can be grown on YMG agar at room temperature (4 g/L d-glucose, 4 g/L yeast extract, 10 g/L malt extract, 18 g/L agar). The culture can be obtained at the Jena Microbial Resource Collection registration number SF:011241.

When the mycelia is wounded by scalpel damage, a yellow pigment appears. These pigments were identified as polyenes by characteristic UV-Vis spectra. The structures of the polyenes were determined by NMR. They were identified as 1) (3Z,5E,7E,9E,11E,13Z,15E,17E)-18-methyl-19-oxoicosa-3,5,7,9,11,13,15,17-octaenoic acid; and 2) (3E,5Z,7E,9E,11E,13E,15Z,17E,19E)-20-methyl-21-oxodocosa-3,5,7,9,11,13,15,17,19-nonaenoic acid. The shorter polyene was found to be highly toxic against Drosophila melanogaster larvae. Feeding experiments with [1-13C]acetate revealed a polyketidic origin, and feeding experiments with l-[methyl-13C]methionine revealed methyl branching likely via S-adenosyl-l-methionine-dependent methyltransferase(s). Additionally, MALDI-TOF imaging at the mycelial wounding site identified these poylenes as localized at the wounded area. Two putative alleles of polyketide genes were identified, referred to as PPS1 and PPS2. QRT-PCR monitoring of PPS1 showed up-regulation of this gene after mycelial wounding. The domain architecture was the following for PPS1: KS, AT, DH, MT, KR, ACP, and thus was classified as a highly-reducing polyketide synthase (HR-PKS). This was the first characterized HR-PKS of basidiomycetic origin and forms its own clade in comparison to ascomycete HR-PKSs, bacterial polyene-forming PKSs, basidiomycete non-reducing PKSs, and ascomycete non-reducing PKSs.[2] This research was highlighted in the media several times.[3]

The fungus also produces fomannoxins and vibralactones. The famonnixins were found to be phytotoxic and the vibralactones were found to be antifungal.[4]

Additionally, a new prenylphenol, termed cloquetin, was identified from the fungus by NMR.[5] Overexpression of two annotated non-reducing polyketide synthases (PKS1 and PKS2) in Aspergillus niger and subsequent characterization of the proteins identified them as orsellinic acid synthases. A prenyltransferase gene, termed BYPB, was overexpressed in E. coli and the subsequent protein was confirmed to add a prenyl group to orsellinic acid.

References

  1. ^ Schwenk, Daniel; Nett, Markus; Dahse, Hans-Martin; Horn, Uwe; Blanchette, Robert A.; Hoffmeister, Dirk (24 November 2014). "Injury-Induced Biosynthesis of Methyl-Branched Polyene Pigments in a White-Rotting Basidiomycete". Journal of Natural Products. 77 (12): 2658–2663. doi:10.1021/np500552a. ISSN 0163-3864. OCLC 5720290006. PMID 25420175.
  2. ^ Brandt et al. (2017) Induced Chemical Defense of a Mushroom by a Double-Bond-Shifting Polyene Synthase. Angewante Chemie.
  3. ^ "Mushrooms Get Defensive: One single enzyme induces the chemical defense of a mushroom against larvae - Wiley News Room – Press Releases, News, Events & Media". newsroom.wiley.com. Retrieved 22 February 2018.
  4. ^ Schwenk et al. (2016) Unexpected Metabolic Versatility in a Combined Fungal Fomannoxin/Vibralactone Biosynthesis. Journal of Natural Products.
  5. ^ Braesel et al. (2016) Biochemical and genetic basis of orsellinic acid biosynthesis and prenylation in a stereaceous basidiomycete. Fungal Genetics and Biology.
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BY1: Brief Summary

provided by wikipedia EN

BY1 is a taxonomically unidentified basidiomycete fungus. ITS sequencing has placed it in the Russulales and is referred to as a stereaceous basidiomycete. Chemotaxonomically supporting its placement in this group, it produces fomannoxins and vibralactones. The fungus' mycelia were isolated from dead aspen in Minnesota, USA. It is presumed to decompose wood by white rot.

The mycelium can be grown on YMG agar at room temperature (4 g/L d-glucose, 4 g/L yeast extract, 10 g/L malt extract, 18 g/L agar). The culture can be obtained at the Jena Microbial Resource Collection registration number SF:011241.

When the mycelia is wounded by scalpel damage, a yellow pigment appears. These pigments were identified as polyenes by characteristic UV-Vis spectra. The structures of the polyenes were determined by NMR. They were identified as 1) (3Z,5E,7E,9E,11E,13Z,15E,17E)-18-methyl-19-oxoicosa-3,5,7,9,11,13,15,17-octaenoic acid; and 2) (3E,5Z,7E,9E,11E,13E,15Z,17E,19E)-20-methyl-21-oxodocosa-3,5,7,9,11,13,15,17,19-nonaenoic acid. The shorter polyene was found to be highly toxic against Drosophila melanogaster larvae. Feeding experiments with [1-13C]acetate revealed a polyketidic origin, and feeding experiments with l-[methyl-13C]methionine revealed methyl branching likely via S-adenosyl-l-methionine-dependent methyltransferase(s). Additionally, MALDI-TOF imaging at the mycelial wounding site identified these poylenes as localized at the wounded area. Two putative alleles of polyketide genes were identified, referred to as PPS1 and PPS2. QRT-PCR monitoring of PPS1 showed up-regulation of this gene after mycelial wounding. The domain architecture was the following for PPS1: KS, AT, DH, MT, KR, ACP, and thus was classified as a highly-reducing polyketide synthase (HR-PKS). This was the first characterized HR-PKS of basidiomycetic origin and forms its own clade in comparison to ascomycete HR-PKSs, bacterial polyene-forming PKSs, basidiomycete non-reducing PKSs, and ascomycete non-reducing PKSs. This research was highlighted in the media several times.

The fungus also produces fomannoxins and vibralactones. The famonnixins were found to be phytotoxic and the vibralactones were found to be antifungal.

Additionally, a new prenylphenol, termed cloquetin, was identified from the fungus by NMR. Overexpression of two annotated non-reducing polyketide synthases (PKS1 and PKS2) in Aspergillus niger and subsequent characterization of the proteins identified them as orsellinic acid synthases. A prenyltransferase gene, termed BYPB, was overexpressed in E. coli and the subsequent protein was confirmed to add a prenyl group to orsellinic acid.

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