Amanita phalloides 184774 223053294 2008-07-02T10:44:50Z Casliber 1381068 cats {{Taxobox | name=Death cap | status= secure | image=Amanita phalloides 1.JPG | image_width=250px | regnum=[[Fungus|Fungi]] | divisio=[[Basidiomycota]] | classis=[[Agaricomycetes]] | subclassis=[[Hymenomycete]]s | ordo=[[Agaricales]] | familia=[[Amanitaceae]] | genus=''[[Amanita]]'' | species='''''A. phalloides''''' | binomial=''Amanita phalloides'' | binomial_authority=([[Sébastien Vaillant|Vaill.]] ex [[Elias Magnus Fries|Fr.]]) [[Johann Heinrich Friedrich Link|Link]] }} {{mycomorphbox | name=Amanita phalloides | whichGills=free | capShape=convex | capShape2=flat | hymeniumType=gills | stipeCharacter=ring and volva | ecologicalType=mycorrhizal | sporePrintColor=white | howEdible=deadly }} '''''Amanita phalloides''''' (generally {{pronEng|æməˈnaɪtə fəˈlɔɪdiːz}}), commonly known as the '''death cap''', is a poisonous [[basidiomycete]] [[fungus]], one of many in the genus ''[[Amanita]]''. Widely distributed across [[Europe]], ''A. phalloides'' [[mycorrhiza|associates]] with various [[deciduous]] trees. In some cases, death cap has been accidentally introduced to new regions with the cultivation of non-native species of [[oak]], [[chestnut]], and [[Pinus|pine]]. The large fruiting bodies (''i.e.'', the [[mushrooms]]) appear in summer and autumn; the [[Pileus (mycology)|caps]] are generally greenish in colour, with a white [[stipe (mycology)|stipe]] and [[Lamella (mycology)|gill]]s. Coincidentally, these toxic mushrooms resemble several edible species (most notably the [[straw mushroom]]) commonly consumed by humans, increasing the risk of accidental [[poison]]ing. ''A. phalloides'' is one of the most poisonous of all known [[toadstool]]s. It has been involved in the majority of human deaths from [[mushroom poisoning]], possibly including the deaths of Roman Emperor [[Claudius]] and [[Charles VI, Holy Roman Emperor|Holy Roman Emperor Charles VI]]. It has been the subject of much research, and many of its biologically active agents have been isolated. The principal toxic constituent is [[alpha-amanitin|α-amanitin]], which damages the liver and kidneys, often fatally. No [[antidote]] is known. ==Taxonomy and naming== The death cap was first described by French botanist [[Sébastien Vaillant]] in 1727, who gave a succinct phrase name "''Fungus phalloides, annulatus, sordide virescens, et patulus''", which is still recognizable as the fungus today.<ref>{{la icon}}{{cite book|last=Vaillant|first=Sébastien|year=1727|title=Botanicon Parisiense|location=Leide & Amsterdam|publisher=J. H. Verbeek and B. Lakeman|oclc=5146641}}</ref> Though the [[scientific name]] ''phalloides'' means "phallus-shaped", it is unclear whether it is named for its resemblance to a literal [[phallus]] or the stinkhorn mushrooms ''[[Phallus (genus)|Phallus]]''. In 1821, [[Elias Magnus Fries]] described it as ''Agaricus phalloides'', but included all white Amanitas within its description.<ref>{{la icon}}{{cite book |last=Fries |first=Elias Magnus |title=Systema Mycologicum I |year=1821 |publisher=Ernesti Mauritii |location=Gryphiswaldiae|oclc=10717479 }}</ref> Finally in 1833, [[Johann Heinrich Friedrich Link]] settled on the name ''Amanita phalloides'',<ref>{{de icon}} Link JHF (1833) ''Grundriss der Kraeuterkunde IV''. Haude und Spenerschen Buchhandlung (S.J. Joseephy), Berlin</ref> after [[Christian Hendrik Persoon|Persoon]] had named it ''Amanita viridis'' thirty years earlier.<ref>{{la icon}}{{cite book |last=Persoon |first=Christian Hendrik |title=Tentamen Dispositionis Methodicae Fungorum |year=1797 |publisher=P.P. Wolf, |location=Lipsiae |oclc=19300194}}</ref><ref>{{la icon}}{{cite book |last=Persoon |first=Christian Hendrik |title=Synopsis Methodica Fungorum |year=1801 |publisher=H. Dietrich |location=GÖttingen|oclc=28329773 }}</ref> Although [[Louis Secretan]]'s use of the name ''Amanita phalloides'' predates Link's, it has been rejected for nomenclatural purposes because Secretan's works did not use [[binomial nomenclature]] consistently;<ref>{{cite journal|last=Donk| first=M.A.| month=June| year=1962| title=On Secretan's Fungus Names| journal=Taxon| volume=11| issue=5| pages=170–173| url=http://links.jstor.org/sici?sici=0040-0262%28196206%2911%3A5%3C170%3AOSFN%3E2.0.CO%3B2-I| doi=10.2307/1216724}}</ref><ref>{{cite journal|last=Demoulin| first=V.| month=November| year=1974| title=Invalidity of Names Published in Secretan's Mycographie Suisse and Some Remarks on the Problem of Publication by Reference| journal=Taxon| volume=23| issue=5/6| pages=836–843| url=http://links.jstor.org/sici?sici=0040-0262%28197411%2923%3A5%2F6%3C836%3AIONPIS%3E2.0.CO%3B2-T| doi=10.2307/1218449}}</ref> some taxonomists have, however, disagreed with this opinion.<ref>{{cite journal|last=Singer| first=Rolf| coauthors=Robert E. Machol| month=June| year=1962| title=Are Secretan's Fungus Names Valid?| journal=Taxon| volume=26| issue=2/3| pages=251–255| url= http://links.jstor.org/sici?sici=0040-0262%28197705%2926%3A2%2F3%3C251%3AASFNV%3E2.0.CO%3B2-G| doi=10.2307/1220563}} </ref><ref>{{cite journal|last=Machol| first=Robert E.| month=August| year=1984| title=Leave the Code Alone| journal=Taxon| volume=33| issue=3| pages=532–533| url=http://links.jstor.org/sici?sici=0040-0262%28198408%2933%3A3%3C532%3ALTCA%3E2.0.CO%3B2-9| doi=10.2307/1221006}}</ref> ''Amanita phalloides'' is the [[type species]] of ''Amanita'' [[Section (botany)|section]] ''Phalloideae'', a group that contains all of the deadly poisonous ''Amanita'' species thus far identified. Most notable of these are the species known as [[destroying angel]]s, namely ''[[Amanita virosa]]'' and ''[[Amanita bisporiga|A. bisporiga]]'' as well as the fool's mushroom ''([[Amanita verna|A. verna]])''. The term "destroying angel" has been applied to ''A. phalloides'' at times, but "death cap" is by far the most common vernacular name used in English. Other common names also listed include "stinking amanita"<ref name = "North67"/> and "deadly amanita".<ref name="benjamin203">Benjamin.p203</ref> A rarely appearing all-white form was initially described ''A. phalloides'' f. ''alba'' by Max Britzelmayr,<ref name=tulloss/><ref name="Jord109">Jordan & Wheeler. p109</ref> though its status has been unclear. It is often found growing amid normally coloured death caps. It has been described, in 2004, as a distinct variety and includes what was termed ''A. verna'' var. ''tarda''.<ref name="Nevpoum04"/> The true ''Amanita verna'' fruits in spring and turns yellow with [[Potassium hydroxide|KOH]] solution, whereas ''A. phalloides'' never does.<ref name=tullossverna>{{Cite web| last=Tulloss| first=Rodham E.| Month=July| Year=2006| url=http://pluto.njcc.com/~ret/amanita/species/verna.html| title= Amanita verna (Bull.: Fr.) Lam.| work=Amanita Studies site|accessdate=2007-05-22}}</ref> ==Description== The death cap has a large and imposing [[epigeous]] (aboveground) [[fruiting body]] (basidiocarp), usually with a [[pileus]] (cap) from 5 to 15&nbsp;cm (2–6&nbsp;[[inch|in]]) across, initially rounded and hemispherical, but flattening with age.<ref name="Bresinsky">{{cite book | author= Bresinsky A, Besl H. | title = A Colour Atlas of Poisonous Fungi | year = 1990 | publisher = Wolfe Publishing | pages=26-9|id = ISBN 0-7234-1576-5}}</ref> The colour of the cap can be pale-, yellowish-, or olive-green, often paler toward the margins and often paler after rain. The cap surface is sticky when wet and easily peeled, a troublesome feature, as that is allegedly a feature of edible fungi.<ref name="Jord99">Jordan & Wheeler. p99</ref> The remains of the [[partial veil]] is seen as a skirtlike, floppy [[annulus (mycology)|annulus]] usually about 1 to 1.5&nbsp;cm (0.4&ndash;0.6&nbsp;in) below the cap. The crowded white [[Lamella (mycology)|lamellae]] (gills) are free. The [[Stipe (mycology)|stipe]] is white with a scattering of grayish-olive scales and is 8 to 15&nbsp;cm (3&ndash;6&nbsp;in) long and 1 to 2&nbsp;cm (3/8&ndash;3/4&nbsp;in) thick, with a swollen, ragged, sac-like white [[volva (mycology)|volva]] (base).<ref name="Bresinsky"/> As the volva, which may be hidden by leaf litter, is a distinctive and diagnostic feature, it is important to remove some debris to check for it.<ref name="Jord108">Jordan & Wheeler. p108</ref> The smell has been described as initially faint and honey-sweet but strengthening over time to become overpowering, sickly-sweet and objectionable.<ref name="Zeitl61">Zeitlmayr. p61</ref> Young specimens first emerge from the ground resembling a white egg covered by a [[universal veil]], which then breaks, leaving the volva as a remnant. The [[spore print]] is white, a common feature of ''Amanita''. The transparent spores are globular to egg-shaped, measure 8–10&nbsp;[[micrometre|μm]] (0.3&ndash;0.4&nbsp;[[Thou (unit of length)|mil]]) long, and stain blue with [[iodine]].<ref name="Zeitl61"/> The gills, on the other hand, are seen to stain pallid lilac or pink with concentrated [[sulfuric acid]].<ref>{{cite book|last=Jordan|first=Michael|year=1995|title=The Encyclopedia of Fungi of Britain and Europe|publisher=David & Charles|pages=198|id=ISBN 0-7153-0129-2}}</ref><ref>{{cite web | title=California Fungi: Amanita phalloides | publisher=MykoWeb.com | url=http://www.mykoweb.com/CAF/species/Amanita_phalloides.html | accessdate=June 01 | accessyear=2007}}</ref> ==Distribution and habitat== [[Image:Amanita phalloïdes.jpg|right|thumb|A young death cap emerging from universal veil ([[France]])]] The death cap is native to [[Europe]], where it is widespread.<ref>{{Cite journal| first=Lene| last=Lange| year=1974| title=The distribution of macromycetes in Europe|journal=Dansk Botanisk Arkiv|volume=30|pages=5–105|issn=0011-6211}}</ref> It is found from the southern coastal regions of [[Scandinavia]] in the north, to [[Ireland]] in the west, east to [[Poland]] and western [[Russia]],<ref name="Nevpoum04">{{Cite book|last=Neville| first=Pierre| coauthors=Serge Poumarat| title=Amaniteae: ''Amanita'', ''Limacella'' and ''Torrendia''| year=2004|series=Fungi Europaei (9)| location=Alassio|Edizioni Candusso|isbn=88-901057-3-9}}</ref> and south throughout the [[Balkans]], in [[Italy]] and [[Spain]], and in [[Morocco]] and [[Algeria]] in north [[Africa]].<ref name="Malber70">{{Cite book|first=Georges|last=Malençon|coauthors=R. Bertault| title=Flore des Champignons Supérieurs du Maroc I| year=1970|location=Rabat|publisher=Faculté des Sciences|series=Travaux de l'Institut scientifique chérifien et de la Faculté des sciences. Série botanique et biologie végétale (32)|oclc=915096}}</ref> There are records from further east into Asia but these have yet to be confirmed as ''A. phalloides''.<ref name="Pringvell06">{{cite journal|last=Pringle| first=Anne| coauthors=Else C. Vellinga|year=2006 |month=July| title=Last chance to know? Using literature to explore the biogeography of and invasion biology of the death cap mushroom ''Amanita phalloides'' (Vaill. Ex Fr. :Fr) Link|journal=Biological Invasions|volume=8|issue=5|pages=1131–1144|doi=10.1007/s10530-005-3804-2}}</ref> It is [[Mycorrhiza#Ectomycorrhizae|ectomycorrhizally]] associated with a number of tree species. In Europe, these include a large number of [[hardwood]] and, less frequently, [[conifer]] species. It appears most commonly under oaks but also under [[beech]]es, [[chestnut]]s, [[Aesculus|horse-chestnuts]], [[birch]]es, [[filbert]]s, [[hornbeam]]s, pines, and [[spruce]]s.<ref name=tulloss>{{Cite web| last=Tulloss| first=Rodham E.| Month=July| Year=2006| url=http://pluto.njcc.com/~ret/amanita/species/phalloid.html| title=(Fr.:Fr.) Link| work=Amanita Studies site|accessdate=2007-05-22}}</ref> In other areas, ''A. phalloides'' may also be associated with these trees or only with some species but not others. In coastal California, for example, ''A. phalloides'' is associated with [[coast live oak]] but not with the various coastal pine species, such as [[Monterey pine]].<ref>{{cite book|last=Arora| first=David| title=Mushrooms demystified : a comprehensive guide to the fleshy fungi| year=1986| location=Berkeley, California| publisher=Ten Speed Press|isbn=0-89815-170-8}}</ref> In countries where it has been introduced it has been restricted to those exotic trees it would associate with in its natural range. There is, however, evidence of ''A. phalloides'' associating with [[Tsuga|hemlock]] and with genera of the [[Myrtaceae]]: ''[[Eucalyptus]]'' in Tanzania<ref name="Peg77">{{cite book| last=Pegler| First=D.N.| year=1977| title=A preliminary agaric flora of East Africa| location:London| publisher= Royal Botanic Gardens, Kew|series= Kew Bulletin Additional Series (6)| isbn=0-11-241101-0}}</ref> and Algeria,<ref name="Malber70"/> and ''[[Leptospermum]]'' and ''[[Kunzea]]'' in New Zealand.<ref name="Rid91">{{cite journal|last=Ridley| first=G.S.|title=The New Zealand Species of ''Amanita'' (Fungi: Agaricales)| year=1991| journal=Australian Systematic Botany| volume=4| issue=2|pages=325–354|doi=10.1071/SB9910325}}</ref><ref name=tulloss/> This suggests the species may have invasive potential.<ref name="Pringvell06"/> By the end of the 19th century, [[Charles Horton Peck]] had reported ''A. phalloides'' in North America.<ref>{{cite book|last=Peck| first=Charles H.| year=1897|title=Annual report of the state botanist|location=Albany|publisher=University of the State of New York|oclc=1185748}}</ref> However, in 1918, samples from the Eastern United States were identified as being a distinct though similar species, ''[[Amanita brunnescens|A. brunnescens]]'', by G. F. Atkinson of [[Cornell University]].<ref name="Litten75">{{cite journal |last=Litten|first= W. |year=1975|month=March |title=The most poisonous mushrooms |journal=[[Scientific American]] |volume=232 |issue=3 |pages=90–101 |pmid=1114308}}</ref> By the 1970s it had become clear that ''A. phalloides'' actually does occur in the United States, apparently having been introduced from [[Europe]] alongside chestnuts, with populations on the West and East Coasts.<ref name="Litten75"/><ref name="benjamin204">Benjamin.p204</ref> A more recent historical review concluded that the East Coast populations were introduced but that the origins of the West Coast population remain unclear, due to the scantness of historical records.<ref name="Pringvell06"/> ''Amanita phalloides'' has been conveyed to new countries across the southern hemisphere with the importation of hardwoods and conifers. Introduced [[oak]]s appear to have been the vector to [[Australia]] and [[South America]]; populations under oaks have been recorded from [[Melbourne]] and [[Canberra]],<ref>{{cite journal |last=Reid |first=D.A. |year=1980 |title=A monograph of the Australian species of ''Amanita'' Pers. ex Hook (Fungi) |journal=[[Australian Journal of Botany]] Supplementary Series |volume=8|pages=1–96}}</ref><ref>{{cite journal |last=Cole| first=F.M. |year=1993| month=June |title=''Amanita phalloides'' in Victoria |journal=Medical Journal of Australia |volume=158 |issue=12 |pages=849–850| pmid=8326898}}</ref> as well as [[Uruguay]].<ref>{{es icon}}{{cite journal|last=Herter|first=W.G.|year=1934| title=La aparición del hongo venenoso ''Amanita phalloides'' en Sudamérica.| journal=Revista Sudamericana de Botánica|volume=1 |pages=111–119}}</ref> It has been recorded under other introduced trees in [[Argentina]]<ref>{{es icon}}{{cite journal| last=Hunzinker| first=A.T.|year=1983| title=''Amanita phalloides'' en las Sierras de Córdoba|journal= Kurtziana| volume=16| pages=157–160|issn=0075-7314}}</ref> and [[Chile]].<ref>{{cite journal| last=Valenzuella| first=E.| coauthors=G. Moreno & M. Jeria| year=1992| title=''Amanita phalloides'' en bosques de ''Pinus radiata'' de la IX Region de Chile: taxonomia, toxinas, metodos de dedection, intoxicacion faloidiana| journal=Boletín Micológico| volume=7| pages=17–21| issn=0716-114X}}</ref> [[Pine]] plantations are associated with the fungus in [[Tanzania]]<ref name="Peg77"/> and [[South Africa]], where it is also found under oaks and [[poplar]]s.<ref>{{cite journal| last=Reid| first=D.A.| coauthors=A. Eicker|year=1991| title=South African fungi: the genus ''Amanita''| journal=Mycological Research|volume=95| issue=1|pages=80–95|issn=0953-7562}}</ref> ==Toxicity== [[Image:Death Cap Mushroom.jpg|thumb|right|Warning sign in [[Canberra]], [[Australia]]]] As the common name suggests, the fungus is highly toxic, and it is responsible for the majority of fatal [[mushroom poisoning]]s worldwide.<ref name="benjamin200">Benjamin.p200</ref> It is estimated that 30&nbsp;[[gram]]s (1&nbsp;oz), or half a cap, of this mushroom is enough to kill a human.<ref name="benjamin211">Benjamin.p211</ref> In 2006, a family of three in [[Poland]] was poisoned, resulting in one death and the two survivors requiring liver transplants.<ref>{{pl icon}} {{cite journal|first=J.|last=Pawlowska|coauthors=Pawlak J, Kamiski A, Hevelke P, Jankowska I, Teisseyre M, Szymczak M, Kaliciiski P, Krawczyk M.| year=2006|title=(''Amanita phalloides'' poisoning as an indication for liver transplantation in three family members.)| journal=Wiadomości Lekarskie| volume=59|issue=1–2|pages=131–4| pmid=16646310}}</ref> Some authorities strongly advise against putting suspected death caps in the same basket with fungi collected for the table and to avoid touching them.<ref name="Jord99"/><ref>{{cite book | author = Carluccio A | year = 2003 | title = The Complete Mushroom Book | publisher = Quadrille | location = London|pages = 224| id = ISBN 1-84400-040-0}}</ref> Furthermore, the toxicity is not [[Thermostability|reduced by cooking]], freezing, or drying. Its biochemistry has been researched intensively for decades.<ref name="Litten75"/> ===Similarity to edible species=== Recent cases highlight the issue of the similarity of ''A. phalloides'' to the [[edible mushroom|edible]] paddy straw mushroom, ''[[Volvariella volvacea]]'', with east- and southeast-Asian immigrants in Australia and the west coast of the United States falling victim. In an episode in [[Oregon]], four members of a [[Korea]]n family required liver transplants.<ref name="benjamin198">Benjamin.p198–199</ref> Of the seven people poisoned in the [[Canberra]] region between 1988 and 1998, three were from [[Laos]].<ref name="trim99">{{cite journal |author=Trim Geoffrey M. ''et al.'' |year=1999 |month=September |title=Poisoning by ''Amanita phalloides'' ("deathcap") mushrooms in the Australian Capital Territory |url=http://www.mja.com.au/public/issues/171_5_060999/trim/trim.html | accessdate=2007-05-22 |journal=[[Medical Journal of Australia]] |volume=171 |issue=5 |pages=247–249 |pmid=10495756}}</ref> This misidentification is a leading cause of mushroom poisoning in the [[United States]]. Novices may mistake juvenile death caps for edible [[puffball]]s<ref name="hall">{{cite book | author= Hall IR, Stephenson SE, Buchanan PK, Yn W, Cole AL. | title = Edible and poisonous mushrooms of the world | year = 2003 | publisher = New Zealand Institute for Crop & Food Research Limited |pages=131-3|id = ISBN 0-4781-0835-4}}</ref> or mature specimens for other edible ''Amanita'' species such as ''[[Amanita lanei]]'', and for this reason some authorities recommend avoiding the collecting of ''Amanita'' species for the table altogether.<ref>{{cite book| last=Phillips| first= Roger| title=Mushrooms and Other Fungi of North America|year= 2005| location=Buffalo| publisher=Firefly books| isbn=1-55407-115-1|pages= p.14}}</ref> The white form of ''A. phalloides'' may be mistaken for edible species of ''[[Agaricus]]'', especially the young fruitbodies whose unexpanded caps conceal the telltale white gills; all mature species of ''Agaricus'' have dark-coloured gills.<ref>{{cite web | last =Heino | first = Lepp| title = Deathcap Mushroom: Amanita phalloides | publisher = Australian National Botanic Gardens | date = 2006-10-09 | url =http://www.anbg.gov.au/fungi/deathcap.html| accessdate = 2007-06-12}}</ref> In Europe, other similarly green-capped species collected by mushroom hunters include various green-hued brittlegills of the genus ''[[Russula]]'' and the formerly popular ''[[Tricholoma flavovirens]]'', now regarded as hazardous owing to a series of restaurant poisonings in [[France]]. Brittlegills, such as ''[[Russula heterophylla]]'', ''[[Russula aeruginea|R. aeruginea]]'', and ''[[Russula virescens|R. virescens]]'', can be distinguished by their brittle flesh and the lack of both volva and ring.<ref name="Zeitl62">Zeitlmayr. p62</ref> Other similar species include ''[[Amanita subjunquillea|A. subjunquillea]]'' in eastern [[Asia]] and ''[[Amanita arocheae|A. arocheae]]'', which ranges from Andean [[Colombia]] north at least as far as central [[Mexico]], both of which are also poisonous. ===Biochemistry=== The species is now known to contain two main groups of toxins, both [[Cyclic compound|multicyclic]] (ring-shaped) [[peptide]]s, spread throughout the mushroom [[Thallus (tissue)|tissue]]: the [[amatoxin]]s and the [[phallotoxin]]s. Another toxin is [[phallolysin]], which has shown some [[haemolysis|hemolytic]] (red blood cell–destroying) activity ''in vitro''. An unrelated compound, [[antamanide]], has also been isolated. Amatoxins consist of at least eight compounds with a similar structure, that of eight amino-acid rings; they were isolated in 1941 by [[Heinrich Otto Wieland|Heinrich O. Wieland]] and Rudolf Hallermayer of the [[Ludwig Maximilians University of Munich|University of Munich]].<ref name="Litten75"/> Of the amatoxins, [[α-amanitin]] is the chief component and along with [[β-amanitin]] is likely responsible for the toxic effects.<ref name="Koppel">{{cite journal |author=Köppel C |title=Clinical symptomatology and management of mushroom poisoning |journal=Toxicon |volume=31 |issue=12 |pages=1513–40 |year=1993 |pmid=8146866 |doi=10.1016/0041-0101(93)90337-I}}</ref><ref name="dart">{{cite book |last=Dart| first= RC |chapter=Mushrooms |title=Medical toxicology |publisher=Williams & Wilkins| location=Philadelphia |year=2004|pages=1719–35| isbn=0-7817-2845-2}}</ref> Their major toxic mechanism is the inhibition of [[RNA polymerase II]], a vital enzyme in the synthesis of [[messenger RNA]] (mRNA), [[microRNA]], and small nuclear RNA ([[snRNA]]). Without mRNA essential [[Protein biosynthesis|protein synthesis]] and hence cell metabolism grind to a halt and the cell dies.<ref name=" Karlson-Stiber">{{cite journal |author=Karlson-Stiber C, Persson H |title=Cytotoxic fungi - an overview |journal=Toxicon |volume=42 |issue=4 |pages=339–49 |year=2003 |pmid=14505933|doi=10.1016/S0041-0101(03)00238-1}}</ref> The [[liver]] is the principal organ affected, as it is the organ which is first encountered after absorption in the gastrointestinal tract, though other organs, especially the [[kidney]]s, are susceptible.<ref name="benjamin217">Benjamin.p217</ref> The RNA polymerase of ''Amanita phalloides'' is insensitive to the effects of amatoxins; as such, the mushroom does not poison itself.<ref name"Horgen">{{cite journal |last==Horgen|first= Paul A.|coauthors= Allan C. Vaisius and Joseph F. Ammirati |title=The insensitivity of mushroom nuclear RNA polymerase activity to inhibition by amatoxins |journal=Archives of Microbiology |volume=118 |issue=3 |pages=317–9 |year=1978 |pmid=567964 |doi=10.1007/BF00429124}}</ref> The phallotoxins consist of at least seven compounds, all of which have seven similar peptide rings. [[Phalloidin]] was isolated in 1937 by [[Feodor Felix Konrad Lynen|Feodor Lynen]], Heinrich Wieland's student and son-in-law, and Ulrich Wieland of the University of Munich. Though phallotoxins are highly toxic to liver cells,<ref>{{cite journal |author=Wieland T, Govindan VM |title=Phallotoxins bind to actins |journal=FEBS Lett. |volume=46 |issue=1 |pages=351–3 |year=1974 |pmid=4429639|doi=10.1016/0014-5793(74)80404-7}}</ref> they have since been found to have little input into the death cap's toxicity as they are not absorbed through the gut.<ref name=" Karlson-Stiber"/> Furthermore, phalloidin is also found in the edible (and sought-after) [[Blusher]] (''Amanita rubescens'').<ref name="Litten75"/> Another group of minor active peptides are the virotoxins, which consist of six similar monocyclic heptapeptides.<ref name = "Vetter"/> Like the phallotoxins they do not exert any acute toxicity after ingestion in humans.<ref name=" Karlson-Stiber"/> ===Symptoms=== Death caps have been reported to taste pleasant.<ref name="Litten75"/><ref name="Clel76">{{cite book | last= Cleland|first= John Burton |origyear=1934 | year=1976| title=Toadstools and mushrooms and other larger fungi of South Australia | publisher=South Australian Government Printer| oclc=15150059 }}</ref> This, coupled with the delay in the appearance of symptoms—during which time internal organs are being severely, sometimes irreparably, damaged—makes it particularly dangerous. Initially, symptoms are [[gastrointestinal]] in nature and include [[Biliary colic|colicky]] abdominal pain, with watery [[diarrhea]] and [[vomiting]] which may lead to [[dehydration]], and, in severe cases, [[hypotension]], [[tachycardia]], [[hypoglycemia]], and [[Acid base physiology|acid-base disturbances]].<ref name="pinson"/><ref name="klein"/> These first symptoms resolve two to three days after the ingestion. A more serious deterioration signifying liver involvement may then occur—[[jaundice]], diarrhea, [[delirium]], [[seizures]], and [[coma]] due to [[fulminant hepatic failure]] and attendant [[hepatic encephalopathy]] caused by the accumulation of normally liver-removed substance in the blood.<ref name = "North67">{{cite book | last=North |first=Pamela Mildred | year=1967 | title=Poisonous plants and fungi in colour| location=London | publisher=Blandford Press|oclc=955264}}</ref> [[Renal failure]] (either secondary to severe [[hepatitis]]<ref>{{cite journal |author=Nicholls DW, Hyne BE, Buchanan P |title=Death cap mushroom poisoning |journal=The New Zealand Medical Journal |volume=108 |issue=1001 |pages=234 |year=1995 |pmid=7603660}}</ref><ref name = "Vetter">{{cite journal|first=János| last=Vetter|year=1998| month=January| title=Toxins of ''Amanita phalloides''| journal=Toxicon| volume=36| issue=1 |pages=13–24|pmid=9604278| doi=10.1016/S0041-0101(97)00074-3}}</ref> or caused by direct toxic renal damage<ref name=" Karlson-Stiber"/>) and [[coagulopathy]] may appear during this stage. Life-threatening complications include increased [[intracranial pressure]], intracranial [[hemorrhage]], [[sepsis]], [[pancreatitis]], [[acute renal failure]], and [[cardiac arrest]].<ref name="pinson"/><ref name="klein"/> Death generally occurs six to sixteen days after the poisoning.<ref>{{cite journal |author=Fineschi V, Di Paolo M, Centini F |title=Histological criteria for diagnosis of amanita phalloides poisoning |journal=J. Forensic Sci. |volume=41 |issue=3 |pages=429–32 |year=1996 |pmid=8656182}}</ref> Up to the mid-20th century, the mortality rate was around 60–70%, but this has greatly improved with advances in medical care. A review of death cap poisoning throughout Europe from 1971 to 1980 found the overall mortality rate to be 22.4% (51.3% in children under ten and 16.5% in those older than ten).<ref name="Floer82">{{de icon}}{{cite journal| last=Floerscheim| first=G.L.| coauthors=O. Weber, P. Tschumi & M. Ulbrich|year=1982|month=August|title=Die klinische knollenblatterpilzvergiftung (''Amanita Phalloides''): prognostische faktoren und therapeutische massnahmen (Clinical death-cap (Amanita phalloides) poisoning: prognostic factors and therapeutic measures.)| journal=Schweizerische medizinische Wochenschrift |volume=112 |issue=34 |pages=1164–1177 |pmid=6291147}}</ref> This has fallen further in more recent surveys to around 10–15%.<ref name="benjamin215">Benjamin.p215</ref> ===Treatment=== Consumption of the death cap is a [[medical emergency]] requiring hospitalization. There are four main categories of therapy for poisoning: preliminary medical care, supportive measures, specific treatments, and [[liver transplantation]].<ref name="Enjalbert">{{cite journal |author=Enjalbert F, Rapior S, Nouguier-Soulé J, Guillon S, Amouroux N, Cabot C |title=Treatment of amatoxin poisoning: 20-year retrospective analysis |journal=Journal of Toxicology - Clinical Toxicology |volume=40 |issue=6 |pages=715–57 |year=2002 |pmid=12475187}}</ref> Preliminary care consists of gastric decontamination with either [[activated carbon]] or [[gastric lavage]]. However, due to the delay between ingestion and the first symptoms of poisoning, it is commonplace for patients to arrive for treatment many hours after ingestion, potentially reducing the efficacy of these interventions.<ref name="Enjalbert"/><ref name="Vesconi">{{cite journal |author=Vesconi S, Langer M, Iapichino G, Costantino D, Busi C, Fiume L |title=Therapy of cytotoxic mushroom intoxication |journal=Critical care medicine |volume=13 |issue=5 |pages=402–6 |year=1985 |pmid=3987318 |doi=10.1097/00003246-198505000-00007}}</ref> Supportive measures are directed towards treating the dehydration which results from fluid loss during the gastrointestinal phase of intoxication and correction of [[metabolic acidosis]], hypoglycemia, [[electrolyte]] imbalances, and impaired coagulation.<ref name="Enjalbert"/> No definitive antidote is available, but some specific treatments have been shown to improve survivability. High-dose continuous intravenous [[Penicillin#Benzylpenicillin .28penicillin G.29|penicillin G]] has been reported to be of benefit, though the exact mechanism is unknown,<ref name="Floer82"/> and trials with [[cephalosporin]]s show promise.<ref name="benjamin227">Benjamin.p227</ref><ref>{{de icon}}{{cite journal|author=Neftel, K. ''et al.''| year=1988| month=January|title=(Are cephalosporins more active than penicillin G in poisoning with the deadly ''Amanita''?)|journal=Schweizerische medizinische Wochenschrift|volume=118| issue=2|pages=49–51|pmid=3278370}}</ref> There is some evidence that intravenous [[silibinin]], an extract from the [[Silybum marianum|blessed milk thistle]] (''Silybum marianum''), may be beneficial in reducing the effects of death cap poisoning. Silibinin prevents the uptake of amatoxins by [[hepatocyte]]s, thereby protecting undamaged hepatic tissue; it also stimulates DNA-dependent RNA polymerases, leading to an increase in RNA synthesis.<ref>{{cite journal |author=Hruby K, Csomos G, Fuhrmann M, Thaler H |title=Chemotherapy of Amanita phalloides poisoning with intravenous silibinin |journal=Human toxicology |volume=2 |issue=2 |pages=183–95 |year=1983 |pmid=6862461}}</ref><ref>{{it icon}} {{cite journal|author=Carducci, R. ''et al.''| year=1996| month=May| title=[[Silibinin and acute poisoning with ''Amanita phalloides'']]|journal=Minerva Anestesiologica|volume=62 |issue=5 |pages=187–93|pmid=8937042}}</ref><ref>{{cite book| last=Jahn| first=W.| coauthors=|year=1980|chapter=Pharmacokinetics of {3H}-methyl-dehydroxymethyl-amanitin in the isolated perfused rat liver, and the influence of several drugs| editor=Helmuth Faulstich, B. Kommerell & Theodore Wieland| title=Amanita toxins and poisoning|publisher=Witzstrock| location=Baden-Baden|pages=80–85|isbn=3-87921-132-9}}</ref> [[Acetylcysteine|N-acetylcysteine]] has shown promise in combination with other therapies.<ref>{{cite journal |author=Montanini S, Sinardi D, Praticò C, Sinardi A, Trimarchi G |title=Use of acetylcysteine as the life-saving antidote in Amanita phalloides (death cap) poisoning. Case report on 11 patients |journal=Arzneimittel-Forschung |volume=49 |issue=12 |pages=1044–7 |year=1999 |pmid=10635453}}</ref> Animal studies indicate the amatoxins deplete hepatic [[glutathione]];<ref>{{cite journal |author=Kawaji A, Sone T, Natsuki R, Isobe M, Takabatake E, Yamaura Y |title=In vitro toxicity test of poisonous mushroom extracts with isolated rat hepatocytes |journal=The Journal of toxicological sciences |volume=15 |issue=3 |pages=145–56 |year=1990 |pmid=2243367}}</ref> N-acetylcysteine serves as a glutathione precursor and may therefore prevent reduced glutathione levels and subsequent liver damage.<ref>{{cite journal |author=Chyka P, Butler A, Holliman B, Herman M |title=Utility of acetylcysteine in treating poisonings and adverse drug reactions |journal=Drug safety |volume=22 |issue=2 |pages=123–48 |year=2000 |pmid=10672895 |doi=10.2165/00002018-200022020-00005}}</ref> None of the antidotes used have undergone prospective, [[randomized clinical trial]]s, and only anecdotal support is available. Silibinin and N-acetylcysteine appear to be the therapies with the most potential benefit.<ref name="Enjalbert"/> Repeated doses of activated carbon may be helpful by absorbing any toxins that are returned to the gastrointestinal tract following [[enterohepatic circulation]].<ref> {{cite journal |author=Busi C, Fiume L, Costantino D, Langer M, Vesconi F |title=Amanita toxins in gastroduodenal fluid of patients poisoned by the mushroom, Amanita phalloides |journal=New England Journal of Medicine |volume=300 |issue=14 |pages=800 |year=1979 |pmid=423916}}</ref> Other methods of enhancing the elimination of the toxins have been trialed; techniques such as [[hemodialysis]],<ref> {{cite journal |author=Sabeel AI, Kurkus J, Lindholm T |title=Intensive hemodialysis and hemoperfusion treatment of Amanita mushroom poisoning |journal=Mycopathologia |volume=131 |issue=2 |pages=107–14 |year=1995 |pmid=8532053 |doi=10.1007/BF01102888}}</ref> [[hemoperfusion]],<ref>{{cite journal |author=Wauters JP, Rossel C, Farquet JJ |title=Amanita phalloides poisoning treated by early charcoal haemoperfusion |journal=British medical journal |volume=2 |issue=6150 |pages=1465 |year=1978 |pmid=719466}}</ref> [[plasmapheresis]],<ref> {{cite journal |author=Jander S, Bischoff J, Woodcock BG |title=Plasmapheresis in the treatment of Amanita phalloides poisoning: II. A review and recommendations |journal=Therapeutic apheresis |volume=4 |issue=4 |pages=308–12 |year=2000 |pmid=10975479| doi=10.1046/j.1526-0968.2000.004004303.x}}</ref> and [[peritoneal dialysis]]<ref> {{cite journal |author=Langer M, Vesconi S, Iapichino G, Costantino D, Radrizzani D |title=The early removal of amatoxins in the treatment of amanita phalloides poisoning |language=German |journal=Klinische Wochenschrift |volume=58 |issue=3 |pages=117–23 |year=1980 |pmid=7366125 |doi=10.1007/BF01477268}}</ref> have occasionally yielded success but overall do not appear to improve outcome.<ref name=" Karlson-Stiber"/> In patients developing liver failure, a liver transplant is often the only option to prevent death. Liver transplants have become a well-established option in amatoxin poisoning.<ref name="klein">{{cite journal|author= Klein AS, Hart J, Brems JJ, Goldstein L, Lewin K, Busuttil RW |year=1989| month=February|title= ''Amanita'' poisoning: treatment and the role of liver transplantation|journal=American Journal of Medicine| volume=86| issue=2| pages=187–93|pmid=2643869|doi=10.1016/0002-9343(89)90267-2}}</ref><ref name="pinson">{{cite journal|author=Pinson CW, Daya MR, Benner KG, Norton RL, Deveney KE, Ascher NL, Roberts JP, Lake JR, Kurkchubasche AG, Ragsdale JW |year=1990|month=May| title=Liver transplantation for severe ''Amanita phalloides'' mushroom poisoning| journal=American Journal of Surgery|volume=159|issue=5|pages=493–9|pmid=2334013|doi=10.1016/S0002-9610(05)81254-1}}</ref><ref>{{cite journal |author=Ganzert M, Felgenhauer N, Zilker T |title=Indication of liver transplantation following amatoxin intoxication |journal=Journal of Hepatology |volume=42 |issue=2 |pages=202–9 |year=2005 |pmid=15664245|doi=10.1016/j.jhep.2004.10.023}}</ref> This is a complicated issue, however, as transplants themselves may have significant [[Complication (medicine)|complications]] and mortality; patients require long-term [[immunosuppression]] to maintain the transplant.<ref name="Enjalbert"/> That being the case, there has been a reassessment of criteria such as onset of symptoms, [[prothrombin time]] (PTT), serum [[bilirubin]], and presence of [[hepatic encephalopathy|encephalopathy]] for determining at what point a transplant becomes necessary for survival.<ref>{{cite journal|last=O'grady| first=John G.| coauthors=Graeme J.M. Alexander, Karen M. Hayllar & Roger Williams|title=Early indicators of prognosis in fulminant hepatic failure| month=august| year=1989| journal=Gastroenterology|volume=97| issue=2| pages=439–445|pmid=2490426}}</ref><ref>{{cite journal| last=Panaro| first=Fabrizio| coauthors=Enzo Andorno, Nicola Morelli, Marco Casaccia, Giuliano Bottino, Ferruccio Ravazzoni, Monica Centanaro, Sara Ornis & Umberto Valente|year=2006| month=April| title=Letter to the editor: Liver transplantation represents the optimal treatment for fulminant hepatic failure from ''Amanita phalloides'' poisoning| journal= Transplant International|volume=19 |issue=4 |pages=344–5| pmid=16573553|doi=10.1111/j.1432-2277.2006.00275.x}}</ref><ref>{{cite journal |author=Escudié L, Francoz C, Vinel JP, Moucari R, Cournot M, Paradis V, Sauvanet A, Belghiti J, Valla D, Bernuau J, Durand F |title=Amanita phalloides poisoning: reassessment of prognostic factors and indications for emergency liver transplantation |journal=J. Hepatol. |volume=46 |issue=3 |pages=466–73 |year=2007 |pmid=17188393 |doi=10.1016/j.jhep.2006.10.013}}</ref> Evidence suggests that, although survival rates have improved with modern medical treatment, in patients with moderate to severe poisoning up to half of those who did recover suffered permanent liver damage.<ref name="benjamin231">Benjamin.p231–232</ref> However, a follow-up study has shown that most survivors recover completely without any [[sequelae]] if treated within 36 hours of mushroom ingestion.<ref>{{cite journal |author=Giannini L, Vannacci A, Missanelli A, Mastroianni R, Mannaioni PF, Moroni F, Masini E |title=Amatoxin poisoning: A 15-year retrospective analysis and follow-up evaluation of 105 patients |journal=Clinical toxicology (Philadelphia, Pa.) |volume=45 |issue=5 |pages=539–42 |year=2007 |pmid=17503263 |doi=10.1080/15563650701365834}}</ref> ==Notable victims== Several historical figures may have died from ''Amanita phalloides'' poisoning (or other similar, toxic ''Amanitas''). These were either accidental poisonings or [[assassination]] plots. Alleged victims of this kind of poisoning include Roman Emperor [[Claudius]], [[Pope Clement VII]], Tsaritsa [[Natalia Naryshkina]], and [[Holy Roman Emperor Charles VI]].<ref name=wasson>{{cite journal|last=Wasson|first=Robert Gordon| year=1972| title=The death of Claudius, or mushrooms for murderers|journal=Botanical Museum Leaflets, Harvard University| volume=23| issue=3|pages=101–128|issn=0006-8098}}</ref> [[R. Gordon Wasson]] recounted the details of these deaths, noting the likelihood of ''Amanita'' poisoning. In the case of Clement VII, the illness that led to his death lasted some five months, making the case clearly inconsistent with amatoxin poisoning. Natalia Naryshkina is said to have consumed a large quantity of pickled mushrooms prior to her death. However, it is unclear whether the mushrooms themselves were poisonous or whether she succumbed to [[food poisoning]].<ref name=wasson/> {{Rquote|right|'''''Ce plat de champignons a changé la destinée de l’Europe.'''''<br /><br /> ("This dish of mushrooms changed the destiny of Europe.")|Voltaire|[http://www.voltaire-integral.com/Html/01/07MEMOIR.html Mémoires].}} Charles VI experienced [[indigestion]] after eating a dish of [[Sautéing|sautéed]] mushrooms. This led to an illness from which he died ten days later — symptomology consistent with amatoxin poisoning. Charles' death led to the [[War of Austrian Succession]]. Noted [[Voltaire]], "this dish of mushrooms changed the destiny of Europe."<ref name="benjamin35">Benjamin.p35</ref><ref name=wasson/> The case of the Claudius poisoning is more complex. It is known that Claudius was very fond of eating ''[[Amanita caesarea|Caesar's mushroom]]''. Following his death, many sources have attributed it to his being fed a meal of death caps instead of Caesar's mushrooms. However, ancient authors such as [[Tacitus]] and [[Suetonius]] are unanimous about there having been poison added to the mushroom dish, rather than the dish having been prepared from poisonous mushrooms. Wasson speculates that the poison used to kill Claudius was derived from death caps, with a fatal dose of [[colocynth]] being administered later during his illness.<ref name="benjamin33">Benjamin.p33–34</ref><ref name=wasson/> ==Footnotes== {{Reflist|3}} ==Sources== *{{cite book |last=Benjamin| first= Denis R. |title=Mushrooms: poisons and panaceas — a handbook for naturalists, mycologists and physicians |publisher=WH Freeman and Company| location=New York |year=1995| isbn=0-7167-2600-9}} *{{cite book | author=Jordan Peter, Wheeler Steven. | year=2001 | title=The Ultimate Mushroom Book | location=London | publisher=Hermes House | isbn = 1-85967-092-X}} *{{cite book|last=Zeitlmayr|first= Linus|year=1976|title=Wild Mushrooms:An Illustrated Handbook|publisher=Garden City Press|location=Hertfordshire|isbn= 0-584-10324-7}} ==External links== {{Spoken Wikipedia|en-Amanita_phalloides.ogg|2007-09-14}} {{commons|Amanita phalloides}} * [http://americanmushrooms.com/deathcap.htm AmericanMushrooms.com - The Death Cap Mushroom ''Amanita phalloides''] * [http://www.mushroomexpert.com/amanita_phalloides.html ''Amanita phalloides'': the death cap] * [http://pluto.njcc.com/~ret/amanita/species/phalloid.html More on this species] * [http://pluto.njcc.com/~ret/amanita/key.dir/phallimb.html Key to species of ''Amanita'' Section ''Phalloidieae'' from North and Central America - ''Amanita'' studies website] * [http://www.anbg.gov.au/fungi/deathcap.html Death cap in Australia - ANBG website] * [http://www.npr.org/templates/story/story.php?storyId=7251327 On the Trail of the Death Cap Mushroom] from American National Public Radio * {{eol|17162994}} {{featured article}} [[Category:Amanita]] [[Category:Fungi of New Zealand]] [[Category:Fungi naturalized in Australia]] [[Category:Fungi of North America]] [[Category:Fungi of Europe]] [[Category:Deadly fungi]] [[Category:Poisonous mushrooms]] {{Link FA|hu}} [[bg:Зелена мухоморка]] [[ca:Farinera borda]] [[cs:Muchomůrka zelená]] [[de:Grüner Knollenblätterpilz]] [[es:Amanita phalloides]] [[eu:Hiltzaile berde]] [[fr:Amanite phalloïde]] [[hr:Zelena pupavka]] [[ilo:Kallugong ti natay (Uong)]] [[is:Grænserkur]] [[it:Amanita phalloides]] [[he:אמנית המוות]] [[lv:Zaļā mušmire]] [[lt:Žalsvoji musmirė]] [[hu:Gyilkos galóca]] [[ms:Cendawan kep kematian]] [[nl:Groene knolamaniet]] [[ja:タマゴテングタケ]] [[no:Grønn fluesopp]] [[pl:Muchomor zielonawy]] [[pt:Amanita phalloides]] [[ru:Поганка бледная]] [[sk:Muchotrávka zelená]] [[sl:Zelena mušnica]] [[sr:Зелена пупавка]] [[fi:Kavalakärpässieni]] [[sv:Lömsk flugsvamp]] [[vi:Nấm tử thần]] [[tr:Amanita phalloides]] [[uk:Мухомор зелений]] [[wa:Tchapea d' macrale]] [[zh:毒鵝膏]]