Sveppir

Skrifað um August 27, 2014 · in Almennt · 1,195 Comments

Áhugi fólks á að matreiða sveppi hefur aukizt talsvert á undanförnum árum. Hér er lerkisveppur í þurrkun. Ljósm. ÁHB.

Áhugi fólks á að matreiða sveppi hefur aukizt talsvert á undanförnum árum. Hér er lerkisveppur í þurrkun. Ljósm. ÁHB.

 

„Eins og þegar er getið, hafa sveppir lengi verið álitnir fremur dularfullar verur, sem stafar af því meðal annars, að mestöll lífsstarfsemi þeirra fer fram niðri í moldinni, hulin sjónum vorum og jafnvel venjulegum rannsóknatækjum.”

 

Helgi Hallgrímsson, 1987: Sveppabaugar og huldurendur

 

„Svöppur k. »sveppur, svampur; knöttur, kúla, …«, svöppur líkl. <*swampu-, sbr. svampur og soppur, sveppur er yngri mynd af svöppur og hefur æxlast af þgf. et. og nf. (og þf.) ft. Líkl. skylt grísku somphós »svampkenndur, gljúpur«.”

 

Ásgeir Bl. Magnússon:

Íslensk orðsifjabók.

1.1 Flokkun

Af gamalli hefð eru sveppir oft taldir til plantna, þó að þeir séu ófrumbjarga. Þeir eru um marga hluti þó ekkert skyldari þörungum og vefplöntum heldur en dýrum. Að vísu eru margir sveppir þráðlaga eins og sumir þörungar og aðeins fáir eru einfrumungar (gersveppir). Engar kvikar frumur koma fyrir í æviferli sveppa og engin bein þróunartengsl virðast vera á milli þeirra og vefplantna.

Steingerðir sveppir eru svo til óþekktir í jarðlögum og því er þróunarsaga þeirra lítið þekkt. Aðeins er unnt að draga ályktanir um uppruna þeirra út frá núlifandi tegun¬dum. Elstu menjar, sem hugsanlega geta verið sveppir, eru um 900 miljón ára, en áreiðanlegar leifar um þá er að finna í 450 til 500 miljón ára gömlum jarðlögum frá ordóvísíum. Ljóst er þó, að fyrstu sveppir hafa verið einfruma heilkjörnungar, en frábrugðnir þeim, sem nú þekkjast. Fyrir um 410 miljón árum hófu lífverur að nema land, og leiddi það til þróunar á gríðarmiklum fjölbreytileika allra lífvera, ekki síður sveppa en allra annarra. Eldri sveppir kunna að hafa lifað í sjó og vatni, en erfiðlega gengur að rekja skyldleika þeirra við núlifandi tegundir. Flest rök hníga að því að fjalla um sveppi sem sjálfstætt ríki – svepparíki.

Hér er sveppum skipt í þrjár fylkingar og innan hinnar síðast nefndu eru taldir þrír flokkar, en að auki eru taldir tveir hópar, sem hafa óvissa stöðu innan flokkunarkerfisins:

OKSVEPPIR (ZYGOMYCOTA)

ASKSVEPPIR (ASCOMYCOTA)

KÓLFSVEPPIR (BASIDIOMYCOTA)

Beðsveppir (Hymenomycetes)

Belgsveppir (Gasteromycetes)

Þelsveppir (Teliomycetes).

[VANBURÐASVEPPIR (FUNGI IMPERFECTI)]

[FLÉTTUR (LICHENES)]

 

Oksveppur.

Oksveppur.

 

Asksveppur.

Asksveppur.

 

Kólfsveppur.

Kólfsveppur.

Allt er óvíst um skyldleika á milli þessara fylkinga. Oksveppir eru sér á báti. Þeir eru samfrumungar, en það merkir, að skil á milli frumna eru horfin, svo að úr verður samrunnin, óslitin heild með mörgum kjörnum. Í asksveppum og kólfsveppum eru skilrúm, götóttir frumuveggir, þar sem umfrymið getur flætt á milli óhindrað. Áður fyrr var álitið, að kólfsveppir væru komnir af asksveppum, en eins má telja víst, að því sé öfugt farið. Þessar þrjár fylkingar voru allar til í upphafi kolatímabilsins fyrir um 300 miljón árum. Eftir steingerðum leifum að dæma eru kólfsveppir miklu mun eldri en asksveppirnir, og styður það kenningu um, að þeir séu forfeður þeirra.

Auk þessara þriggja fylkinga er einn hópur sveppa einstakur og af sumum talinn sérstakur flokkur innan asksveppa eða jafnvel einstök fylking. Það eru vanburðasveppir (Fungi imperfecti). Til þeirra teljast tegundir, sem hefur ekki enn tekizt að fella að ofan nefndri flokkun; sumir nefna þá »ófullkomna sveppi«. Áður var á það minnzt, að fléttur (skófir) eru sveppir og þörungar eða blágerlar í sambýli. Svo náin tengsl eru á milli lífveranna tveggja, að litið er á fléttur sem sérstakar tegundir og er oftast fjallað um þær undir sveppum, því að sveppurinn er umfangsmeiri en hinn aðilinn.

Nú hefur verið lýst um 100’000 tegundum af sveppum. Ekki er ósennilegt, að álíka margar tegundir til viðbótar leynist enn óskráðar. Af fléttum eru þekktar um 25’000 tegundir. – Til sveppa eru líka oft taldar ýmsar aðrar ófrumbjarga lífverur, sem hér eru taldar til frumvera, þar sem engin augljós tengsl eru á milli þeirra og eiginlegra sveppa. Af gamalli hefð bera þeir enn sveppanafn.

 

 

1.2 Bygging sveppa

Sveppir eru fyrst og fremst landlífverur. Flestir eru fjölfrumungar, þó að til séu nokkrar einfruma tegundir (gersveppir). Í raun er hver sveppur gerður úr þráðum, sem nefnast ímur (hyphae). Þræðirnir eru hárfínir og sjaldnast sjáanlegir, nema þeir vaxi margir saman á yfirborði hluta og kallast þá mygla. Knippi af þeim kallast sveppþal (mýsli, mycelium). Vöxtur ímu fer fram í broddinum en myndun prótína fer fram alls staðar í þræðinum og flytjast prótínin til innan ímunnar fyrir tilstilli frymisstrauma, sem eru ríkulegir í þræðinum. Það, sem flestum er tamt að kalla sveppi, svo nefndir stórsveppir, eru í raun knippi af slíkum þráðum, sem eru þétt ofnir saman.

Bæði vegna skjóts vaxtar og þráðlögunar sinnar hafa sveppir mikla sérstöðu með tilliti til umhverfisins borið saman við aðrar lífverur. Hlutfallið á milli yfirborðs og rúmmáls hjá sveppum er mjög hátt. Það merkir, að þeir eru í mikilli snertingu við umhverfið, ef svo má að orði komast. Enginn hluti sveppa er meira en örfáa míkrómetra frá umhverfi; það, sem skilur á milli, er þunnur frumuveggur og næfurþunn frumuhimna. Vegna þess hve sveppþalið er víðfeðmt getur sveppur haft mikil áhrif á umhverfið, meðal annars með því að binda saman agnir í jarðvegi. Oft og tíðum gróa ímur saman, meira að segja þær, sem vaxið hafa frá tveimur gróum. Á þennan hátt getur myndazt stór samfelld breiða af sveppþráðum.

Fyrir því hve náin tengsl eru á milli sveppa og umhverfis, verða allir hlutir sveppsins að vera virkir. Í sumum plöntum er hluti vefja gerður úr dauðum frumum, svo sem viðarvefur, en slíkt er með öllu óþekkt meðal sveppa. Enzým og önnur efni, sem sveppir seyta út í umhverfið hafa áhrif þegar í stað og gegna mikilvægu hlutverki fyrir sveppina sjálfa.

Allir sveppir eru klæddir með frumuvegg. Í vefplöntum og mörgum frumverum er hann gerður úr beðmi (sellulósa), ásamt öðrum sameindum eins og hálfbeðmi (hemísellulósa) og pektíni. Í sveppum hins vegar er frumuveggur úr annarri fjölsykru – kítíni – sem er líka að finna í skeljum og ytra stoðkerfi liðfætlna, svo sem skordýrum, kóngulóm og krabbadýrum. Kítínið hefur þann kost, að það er miklu þolnara gagnvart niðurbroti örvera heldur en beðmið.

Sveppir hafa líffræðilega sérstöðu um marga hluti, sem enn er ekki að fullu þekkt. Eitt hið markverðasta er hvernig kjarnaskiptingu er öðru vísi háttað en í vefplöntum, vefdýrum og flestum frumverum. Á meðan á henni stendur hverfur kjarnahjúpur ekki. Spóluþræðirnir myndast inni í kjarnanum en deilikorn eru engin þekkt. Á síðasti stigi skiptingar herpist kjarnahjúpurinn saman á milli tveggja dótturkjarna og verður að tveimur. Þessi sérstaða undirstrikar enn frekar, að sveppir eru ekki nátengdir öðrum heilkjörnungum. Þá er vert að geta þess, að á engu æviskeiði þeirra koma fyrir frumur búnar svipum eða bifhárum. Meðal annars vegna þessa eru sveppir taldir til sérstaks ríkis.

Kynæxlun sveppa er afar flókin. Æxlunarfærin eru skilin frá ímu með heilum vegg, og myndast kynfrumur, sem allar eru jafnstórar og eins að gerð (jafnfrjóvgun, isogamy), í sérstökum kynhirzlum (gametangia). Nöfn á fylkingum sveppa eru dregin af þessum hirzlum; okfrumuhirzla, askur og kólfur. Við samruna tveggja kynfrumna, það er frjóvgun, verður til tvílitna okfruma, og skiptir hún sér þegar í stað meiósu-frumuskiptingu og verður að fjórum einlitna meiósugróum.

Við kynlausa æxlun myndast gróin í svo nefndum gróhirzlum (sporangia) hjá oksveppum en kóníðuhirzlum (sprotagróhirzlum) hjá ask- og kólfsveppum. Gróin, sem verða til við kynlausa æxlun, myndast við mítósu og eru því réttnefnd mítósugró.

Upp af einni grófrumu getur sprottið fullvaxin lífvera. Gró sveppa geta ekki hreyft sig sjálf, eins og áður hefur komið fram, og dreifast þau aðallega með vindi. Mörg þeirra eru mjög lítil og geta því svifið æði lengi í lofti og stuðlar það að víðáttumikilli útbreiðslu. Sum gró festast auðveldlega við ýmis dýr og dreifast með þeim. Nokkrar sveppategundir (skotsveppir, Pilobolus sp.) skjóta gróunum hátt í loft upp.

 

Cladosporium herbarum.

Cladosporium herbarum.

1.3 Lífshættir sveppa

Sveppir geta vaxið við afar mismunandi aðstæður. Meðal annars getur tegundin Cladosporium herbarum vaxið við –10°C og skemmt kjöt, þó að það sé geymt í kæli. Þá vex ein tegund innan ættkvíslarinnar Chaetomium lang best við 50°C og þrífst meira að segja dável við 60°C.

Allir sveppir eru ófrumbjarga. Þeir afla sér fæðu ýmist sem rotverur eða sníklar. Lífshættir þeirra eru því einkum með tvennu móti, rotlífi og sníkjulífi; að auki lifa margir í sambýli við aðrar lífverur. Þeir soga til sín fæðu eftir að hún hefur að hluta til verið brotin niður áður fyrir tilstilli enzýma, sem sveppirnir gefa frá sér. Margir rotsveppir hafa einkar sérhæfðar ímur, rætlinga, sem festa þá tryggilega við næringarefnið, sem þeir lifa á. Sama er að segja um sníkjusveppi, sérhæfðar ímur, þjófangar (haustorium), draga næringu beint úr frumum hýsilsins. Eftirtektarvert er, að sveppir leggjast miklu heldur á plöntur en dýr; öfugt við gerla.

Sumir sveppir, einkum gersveppir, geta losað orku við gerjun, eins og við myndun alkóhóls úr glúkósa. Helzta forðafjölsykra í sveppum er glýkógen eins og í dýrum og gerlum, en lípíð (fituefni) kunna einnig að safnast fyrir í nokkrum þeirra.

 

 

1.3.1 Rotlífi

Niðurbrot lífrænna efna – það er rotnun dauðra lífvera – er ekki síður mikilvægt en fæðing nýs lífs. Án dauða og rotnunar væri ekkert líf til. Sérhver sameind, sem er að finna í lífverum um stundasakir, er að meira eða minna leyti í sífelldri hringrás. Sveppir eru, ásamt ófrumbjarga gerlum og nokkrum ófrumbjarga lífverum öðrum, mikilvirkustu rotverur hér á jörðu. Við rotnun lífrænna efna losnar koltvíoxíð (CO2) á nýjan leik út í andrúmsloftið og nitursambönd og önnur steinefni blandast öðrum jarðefnum og halda áfram óslitinni hringrás sinni í öðrum plöntum eða dýrum.

Rotsveppir valda oft talsverðu tjóni á mannvirkjum. Í þeim myndast enzým, sem þeir seyta út í umhverfið og brjóta niður lífræn efni, hvort heldur það er fallið tré í birkiskógi eða girðingarstaur úr rekaviði. Hér er því um nokkurs konar útvortis meltingu að ræða. Rotnunin er mest í hitabeltislöndum vegna þess, að hiti og raki örva mjög vöxt sveppa. Í seinni heimsstyrjöldinni var talið að minna en helmingur af öllum aðsendum aðföngum í hitabeltinu hafi verið nothæfur vegna þess að sveppir lögðust á föt, málningu, pappaumbúðir, hjólbarða, leður, vaxefni, þotueldsneyti, einangrun á rafmagnsvírum, filmur og meira að segja hlífðarhúð á linsum í ýmsum sjóntækjum. Í raun má segja, að flest allt varð meira eða minna fyrir barðinu á rotsveppum. Í tempraða beltinu valda sveppir líka verulegu tjóni á matföngum. Þeir vaxa á brauði, grænmeti, ávöxtum, mjöli og mörgu öðru. Sveppirnir rýra verulega næringargildi fæðunnar auk þess sem þeir spilla bragði. Þá eru margir sveppir eitraðir og þarf mjög lítinn skammt af eitri sumra þeirra til þess að illa fari.

1.3.2 Sníkjulífi

Fjölmargir sveppir leggjast á lifandi lífverur og taka til sín næringu úr þeim. Nokkrar tegundir valda um leið ýmis konar, misalvarlegum sýkingum; talið er að um 5000 sveppategundir geti skaðað nytjaplöntur og garðplöntur og einnig leggjast þær á villtar tegundir. Sumar tegundir lifa í viðarvef lifandi trjáa og skerða viðarframleiðslu hvers árs um óheyrilega marga teningsmetra.

Sníkjusveppir geta verið svo aðgangsharðir gagnvart hýsli sínum, að hann drepst skjótlega. Sýnu fleiri eru þó dæmin um það, að sveppurinn lifi í áraraðir á hýsli án augljósrar kopunar; þess eru meira að segja dæmi, að sníkjusveppir valdi auknum vexti. Af nokkrum plöntusjúkdómum, sem sveppir valda, má nefna reyniátu á reyniviði og birkiryð á birkitrjám. Sjúkdómarnir hafa valdið nokkru tjóni hér á landi.

Á milli sveppa og annarra lífvera eru margs konar önnur tengsl, eða aðrar gerðir samlífis eins og meðal gerla og annarra lífvera. Sem dæmi um samhjálp má nefna, að á rótum margra vefplantna (allt að 80%) lifa sveppir og mynda svo nefnda svepprót. Þá eru fléttur annað dæmi um samhjálp á milli sveppa og þörunga eða blágerla.

1.4 Oksveppir

Um 600 tegundir hafa verið skilgreindar innan fylkingarinnar oksveppir. Flestir þeirra lifa á rotnandi plöntum og dýrum í jarðvegi. Sumir eru sníklar á plöntum, skordýrum og smávöxnum jarðvegsdýrum og örfáir geta valdið alvarlegum sýkingum í mönnum og húsdýrum. Flestar tegundir innan fylkingarinnar eru samfrumungar, það er ímurnar eru fjölkjarna, því að frumuveggir eru engir á milli frumna.

 

Brauðmyglusveppur (Rhizopus stolonifer).

Brauðmyglusveppur (Rhizopus stolonifer).

Mygla á brauði heyrir til þessum hópi, enda er »svarti brauðmyglusveppur« (Rhizopus stolonifer) einn þekktasti oksveppurinn. Hann lítur út eins og svartur ullarflóki á rökum, sykruríkum fæðutegundum. Sveppurinn nærist líka á ávöxtum og grænmeti og skemmir þau fljótt. Sveppþalið er úr þrenns konar einlitna sveppþráðum, ímum. Meginhluti þalsins er fljótsprottinn, vex á ætinu og dregur til sín mest af næringu. Út frá þessum ímum vaxa nokkurs konar renglur, sem mynda rætlinga alls staðar þar, sem broddur renglunnar snertir ætið.

Sveppurinn fjölgar sér mjög ört kynlaust. Beint upp af rætlingunum vaxa gróstilkar með gróhirzlum á endum. Hver gróhirzla byrjar sem lítill hnúður, sem þenst út, og streyma frumukjarnar inn í hana. Að skömmum tíma liðnum afmarkast gróhirzlan af nýmynduðum frumuvegg. Umfrymið innan hirzlunnar hólfast niður og veggur tekur að vaxa utan um hvert gró. Gróhirzlurnar verða svartar við þroskun og gefa sveppnum hinn einkennandi lit. Hvert gró, sem losnar, getur orðið að nýju þali. Þessi gró verða til við venjulega frumuskiptingu, mítósu og kallast því mítósugró.

Fræðiheitið Zygomycota (gríska zygon: ok, helsi) er dregið af einu megineinkenni fylkingarinnar, sérstakri kynhirzlu, okfrumuhirzlu, sem myndast að loknum samruna tveggja fjölkjarna kynhirzlna. Kynhirzlurnar þurfa að vera á tveimur ólíkum ættstofnum, sem jafnan eru táknaðir með + og –. Inni í okfrumuhirzlunni verður til ein eða margar tvílitna okfrumur. Okfrumuhirzlan er dvalastig, en við réttar aðstæður verður meiósa í okfrumunum, þær taka að spíra og mynda gróhirzlur með aragrúa gróa. Þessi gró hafa orðið til við meiósu og kallast því meiósugró.

 

Æviferill brauðmyglusvepps (Rhizopus stolonifer).

Æviferill brauðmyglusvepps (Rhizopus stolonifer).

Okfrumuhirzlan hefur á stundum verið kölluð okgró, en þar sem hún hefur að geyma marga tvílitna kjarna (okfrumukjarna) er það ekki alls kostar rétt

 

 

1.5 Asksveppir

1.5.1 Einkenni asksveppa

Um 30’000 tegundum hefur verið lýst innan fylkingar asksveppa. Asksveppir hafa verið nefndir ýmsum nöfnum öðrum eins og sekksveppir, pokagrósveppir og eskisveppir. Margir þeirra eru algengir og sumir hafa hagnýta þýðingu. Margar skrautlegar mygluskánir, rauðar, blágrænar og brúnar, sem skemma matvæli, heyra til asksveppa, svo og hinn laxaliti kimbasveppur, Neurospora sp., sem mikið hefur verið rannsakaður í sambandi við athuganir í erfðafræði. – Hér á landi hefur verið getið um 590 tegundir asksveppa. Flestir vaxa á plöntum, dauðum og lifandi, rekaviði og taði.

Asksveppir eru allir þráðlaga á meðan þeir eru í vexti, nema gersveppir. Ímurnar eru venjulega með þverveggjum, sem eru götóttir, svo að umfrymið og kjarnarnir geta streymt um allan þráðinn. Hver fruma er ýmist með einum eða fleirum kjörnum. Á nokkrum tegundum er þalið einsleitt (homothallic) og geta hvaða þræðir sem er runnið saman, en aðrar tegundir hafa misleitt þal (heterothallic), og geta aðeins + og – ímur runnið saman.

 

Skarlatsdiskur (Aleuria aurantia (Fr.) Fuckel). Stjórnarsandi, 6. September 1997. Ljósm. ÁHB.

Skarlatsdiskur (Aleuria aurantia (Fr.) Fuckel). Stjórnarsandi, 6. september 1997. Ljósm. ÁHB.

 

1.5.2 Kynlaus æxlun

Flestir asksveppir fjölga sér kynlaust með sérstakri gerð af mítósugróum og stuðla þau aðallega að útbreiðslu sveppsins. Þau nefnast kóníðugró eða sprotagró (conidia, dregið af grísku orði, sem merkir fínt ryk) og eru oftast fjölkjarna. Kóníðugróin verða til úr sérstökum móðurfrumum á kóníðusprotum. Sprotarnir greinast oft sérkennilega og geta vafist saman í sérlegar myndanir. Margar tegundir fjölga sér eingöngu með kóníðugróum.

 

Kóníðugró á pensillínsveppi (Penicillium sp.) og bursthýi (Aspergillus sp.)

Kóníðugró á pensillínsveppi (Penicillium sp.) og bursthýi (Aspergillus sp.)

1.5.3 Kynæxlun

Kynæxlun felur ætíð í sér myndun á aski (ascus), sekklaga hirzlu með einlitna askgróum. Bæði askurinn og askgróin eru einstæðar myndanir, sem greina asksveppi frá öllum öðrum sveppum.

Askarnir myndast úr sérstökum frumum. Inni í þeim verða til að loknu flóknu ferli tveir kjarnar eftir að tveir óskyldir sveppþræðir hafa tengzt saman. Þeir renna að lokum saman og mynda okfrumu. Stuttu eftir kjarnasamrunann byrjar að togna úr askinum, og tvílitna kjarni skiptir sér meiósu. Að henni lokinni verða til fjórir einlitna kjarnar. Venjulega fylgir mítósa strax á eftir, svo að til verða átta kjarnar, annars fjórir í hverjum aski. Þessir fjórir eða átta kjarnar innan frumunnar hólfast síðan af, þroskast og verða að fullmynduðum askgróum.

Askar með gróum (mikil stækkun).

Askar með gróum (mikil stækkun).

 

Í flestum asksveppum þrútnar askurinn við þroskun, en að lokum brestur hann og spýtir askgróunum út í loftið. Venjulega þeytast gróin aðeins um tvo sentímetra en þó eru til dæmi um, að þau þjóti allt að 30 sentímetra.

 

Askar verða til inni í sérstökum askhirzlum, og eru margar þeirra vel sýnilegar. Í hverri askhirzlu eru einnig ímur, sem urðu ekki að öskum og nefnast þær geldímur. Askhirzlur eru einkum af tveimur gerðum, annars vegar meira eða minna opnar og disklaga (apothecium) eða flöskulaga með opi efst, þar sem askgróunum er skotið út (perithecium). Askhirzlur hafa aldrei fundizt hjá allmörgum tegundum, og þær eru því taldar til vanburðasveppa.

 

Tvær aðalgerðir af askhirzlum. Í þeim þroskast askarnir.

Tvær aðalgerðir af askhirzlum. Í þeim þroskast askarnir.

1.5.4 Einfruma asksveppir: Gersveppir

Flestir gersveppir eru einfrumungar. Þeim fjölgar venjulega kynlaust, við mítósu, og lítur út sem afkvæmisfruman vaxi út úr móðurfrumunni (knappskot) en sjaldnar með gróum. Kynæxlun verður við það, að tvær frumur eða tvö askgró sameinast og mynda okfrumu. Okfruman kemur í stað asks og innan hennar verður meiósa, svo að fjórir einlitna kjarnar myndast. Þá getur og mítósa fylgt í kjölfarið og verða kjarnarnir þá átta samtals. Frumuveggir myndast um kjarnana, sem þá verða að askgróum, askurinn rifnar eða grotnar og gróin rjúka út í veður og vind.

Mikilvægi gersveppa er ekki sízt fólgið í því, að þeir geta valdið gerjun, það er brotið niður sykur og myndað vínanda (etanól) og koldíoxíð. Víngerðarmenn nota gersveppi til að framleiða vínanda, bakarar sækjast eftir koltvíoxíðinu og ölgerðarmennirnir eftir hvoru tveggja. Fram hafa komið margir ólíkir stofnar af gersveppum og hin síðari ár hafa þeir verið mikið notaðir í margvíslegum líftækniiðnaði.

Saccharomyces cerevisiae.

Saccharomyces cerevisiae.

Kunnasti gersveppurinn við þessa framleiðslu er Saccharomyces cerevisiae, sem ranglega hefur verið nefndur ölgerill á íslensku. Af honum eru til margir stofnar. Tegundin er líka mjög mikið notuð við allskyns rannsóknir í erfða- og frumufræði. Korndrjóli (melaskítur; Claviceps purpurea) er annar gersveppur, sem myndar hörð og íbjúg »berserkjakorn« í axi grastegunda. Hann sýkir ýmsar korntegundir og eitrið í honum veldur svima, sjóndepru og samdrætti æða (drep í fingrum og tám). Nokkur lyf hafa verið unnin úr korndrjóla, meðal annars notað í fæðingarlækningum og mígrenulyfið ergotamín, enda nefnist korndrjóli ergot á ensku.

Þótt flestir einfruma sveppir séu asksveppir eru örfáir af ættkvíslum kólfsveppa, t.d. Cryptococcus neoformans og Candida albicans, sem báðir valda sjúkdómum í mönnum. Ljóst þykir, að hinir einföldu einfrumungar hafa orðið til af flóknari gerðum, en sökum þess, hve fábrotnir þeir eru nú er örðugt að greina forfeður þeirra. Þróun gersveppa úr fjölfruma sveppþali hefur gerst á löngum tíma og oftar en einu sinni. Nú eru þekktar um 40 ættkvíslir gersveppa og heyra til þeirra um 350 tegundir. Þeir lifa bæði á landi og í vatni við ærið margbreytilegar aðstæður.

 

 

1.6 Kólfsveppir

Til þessarar fylkingar, kólfsveppa, heyra um 25’000 tegundir; ekki einungis þær, sem almenningur kallar sveppi, svo sem ullblekill, furusúlungur og lerkisveppur, heldur líka gorkúlur, ryð- og sótsveppir, sem sníkja á plöntum. Enda þótt kólfsveppir séu allvel þekktir, er talið næsta víst, að enn séu til margar óþekktar tegundir. Fjölbreytnin er mjög mikil innan fylkingarinnar. Víða um heim eru fjölmenn áhugamannafélög um gagnsemi og nýtingu sveppa.

Gulbroddi (Hydrum repandum).

Gulbroddi (Hydrum repandum).

1.1.1 Æxlun

Kólfsveppir greinast frá öðrum sveppum á því, að þeir mynda kólfgró (basíðugró) á kylfulaga gróberum, sem nefnast kólfar (basidium). Venjuleg endafruma á ímu þroskast og verður að kólfi. Í frumunni eru tveir einlitna kjarnar. Þeir renna saman og verða að einum tvílitna kjarna. Sá kjarni skiptir sér fljótt meiósu, svo að til verða fjórir einlitna kjarnar. Að þessu loknu myndast fjórir smáir nabbar ofan á kólfi, það tognar smám saman úr þeim og þeir verða að stuttum stilkum, með lítið eitt útblásnum belgjum á endum. Nú leita kjarnarnir fjórir hver í sinn stilk og út í belgina, sem verða að kólfgróum. Þegar gróin ná fullum þroska falla þau af.

Vöxtur kólfa.

Vöxtur kólfa.

Kólfarnir þurfa ljós til þess að þroskast. Sveppþalið, sem kólfarnir sitja á, tekur ýmsum breytingum við þroskunina og sveppþræðirnir sérhæfast til margvíslegra starfa. Sveppþalið með kólfunum raðar sér saman og myndar gróbeð. Inn á milli kólfanna eru geldímur og fingurlaga frumur, sem þumlur nefnast. Talið er, að þessir þræðir séu ófrjóir, ummyndaðir kólfar.

 

1.1.2 Flokkar kólfsveppa

Kólfsveppum má skipta í þrjá flokka eftir gerð gróbeðjar,

a) beðsveppi,
b) belgsveppi
c) þelsveppi.

 

Til beðsveppa heyra flestir sveppir, sem almenningur þekkir hvað best, bæði ætir og eitraðir. Þeir eru oft nefndir hattsveppir. Kólfgróin myndast utan á gróbeðinum gagnstætt því, sem gerist hjá belgsveppum, þar sem þau myndast inni í honum.

Beðsveppir minna einatt á staf með hatti eða hettu á (hattsveppir). Á neðra borði hattsins eru fanir, sem liggja eins og geislar út frá stafnum og minna á hnífsblöð. Þær eru alþaktar kólfum og fá á sig lit gróa, þegar þau eru full þroskuð. Fanirnar eru á stundum greindar eins og gaffall en tengdar saman með þverveggjum, svo að þær líta ekki út eins og hnífsblöð heldur þéttstæð hólf eða rör. Nafnið pípusveppir er af því dregið. Til þeirra teljast helstu matsveppir hér á landi eins og kóngssveppur, kúalubbi, lerki- og furusúlungur.

Fólki er ráðlagt að halda sig við pípusveppi, langi það til þess að leggja sér sveppi til munns, en þekkir lítið til sveppa.

Þegar hattsveppir teygja sig upp úr mold eru þeir oft klæddir innri eða ytri himnum (hulum) eða hvoru tveggja, svo að þeir líkjast belgsveppum. Ytri himnan, heildarhula eða reifar, klæðir allan sveppinn, en innri himnan, fanhula, umlykur hattbörðin og nær niður á staf. Á fullþroskuðum sveppi má oftast sjá leifar af þessum hulum og hafa þær sérstök nöfn, enda mjög mikilsverð greiningareinkenni (sjá mynd). Leifar heildarhulu á hatti nefnast hattflyksur en skeið á stafnum. Leifar fanhulu á staf nefnast kragi (eða hringur) og hengsli á hattbarði. Þeir hattsveppir, sem hafa engar himnur um sig eru sagðir berir.

 

Langskurður af ungum og þroskuðum hattsveppi.. 1. heildarhula (myndar hattflyksur og skeið), 2. fanhula (myndar hengsli og kraga), 3. fanir, 4. stafur, 5. hattur

Langskurður af ungum og þroskuðum hattsveppi.. 1. heildarhula (myndar hattflyksur og skeið), 2. fanhula (myndar hengsli og kraga), 3. fanir, 4. stafur, 5. hattur.

Nokkrar tegundir beðsveppa skortir hina dæmigerðu lögun og verða því trauðla nefndir hattsveppir. Þeir eru margbreytilegir að lögun og eru nefndir vanfönungar eða hattleysingjar.

Í belgsveppum þroskast kólfgró í lokuðum gróhirzlum, ýmist í eða ofan á jarðvegi. Belgurinn utan um kólfgróhirzluna er hnöttóttur, egg-, peru- eða bikarlaga og nefnist byrða og er úr tveimur lögum.

Hér á landi vaxa um þrjátíu tegundir belgsveppa. Kunnastur þeirra er sortukúla eða kerlingareldur (Bovista nigriscens), sem þykir ágætur matsveppur. Innan í belgnum, þar sem gróin þroskast, er sveppþalið, gyrjan, oft ostkennt (merarostur) í byrjun, þá verður það gulbrúnt og að lokum þurrt eins og ull og rjúka gróin út í veður og vind við minnstu snertingu (kerlingareldur). Nafnið fýsisveppir er að því dregið, að gróin rjúka út um kringlótt op, sem myndast á þroskuðum belgnum við minnstu snertingu.

„Fýsisveppur. Þegar þessi sveppur er orðinn þurr, kallast hann kerlingareldur og græðisveppur. Þetta síðasta nafn hefur hann fengið af því hann græðir vel opin sár, sem ei eru djúp, og stillir hann blóðrás úr þeim, en hvorttveggja verkar helst rykið sem í honum er; það er fræ [gró] hans. Það ryk er líka gott við ill fótasár. Finnar gefa þetta ryk kálfum við lífsýki, og blanda því í mjólk.
Ol. Borrichius ráðleggur að skera þennan svepp í tvennt og leggja hann yfir æð, sem skorin er í sundur til að stilla blóðrásina.
Það er almenn sögn, að ryk úr þessum sveppi þurrum blindi mann, ef það kemur í augu hans.
Að hann sé eitur, ef hann kemur í maga manns, kann eg ekki fyrir víst segja, en ráðlegra mun að forðast hans nautn á meðan reynslan hefir ei helgað hann til manneldis.”

Björn Halldórsson, 1783: Grasnytjar.

Í hreiðursveppum geymast kólfgróin í smáum kúlulaga hylkjum, byrðlingum, í botni byrðunnar, sem opnast með stóru opi að ofan. Byrðan líkist hreiðri og byrðlingum svipar til eggja í hreiðri; er nafnið af því dregið.

 

Til þelsveppa teljast tveir stórir ættbálkar, ryðsveppir og sótsveppir. Þeir mynda engan eða mjög lítinn gróbeð og greinast á því frá öðrum kólfsveppum. Kólfar þeirra eru oft skiptir í fjórar frumur, og því eru þeir nefndir brotkólfungar. Þessir sveppir eru yfirleitt sníkjusveppir á plöntum og geta valdið stórtjóni.

Ryðsveppir eru oft á neðra borði blaða og mynda þar ryðbrúnar eða gulbrúnar skellur. Eins og nafnið bendir til mynda sótsveppir svartar, sótkenndar skellur á ýmsum plöntum.

Bláberjalyngsroði. Sníkjusveppur úr hópi kólfsveppa veldur sérkennilegum lit á árssprotum á bláberjalyngi en líka ofvexti, svo að blöðin verða oft stærri en vanalega. Er algengur víða um land en virðist ekki valda varanlegum skemmdum.

Bláberjalyngsroði. Sníkjusveppur úr hópi kólfsveppa veldur sérkennilegum lit á árssprotum á bláberjalyngi en líka ofvexti, svo að blöðin verða oft stærri en vanalega. Er algengur víða um land en virðist ekki valda varanlegum skemmdum.

 

1.7 Svepprót og nornabaugar

Margar tegundir sveppa (einkum hattsveppa) eru háðar trjám. Þær þurfa þó ekki að vera sníkjuverur, heldur mynda þær svo kallaða svepprót. Meðal þessara tegunda myndar sveppþalið hulstur umhverfis ungar trjárætur og smýgur inn á milli frumna í ystu lögum rótar (útræn (ektotrof) svepprót). Sumir sveppir skjóta þjóföngum inn í frumur (ektendotrof svepprót). Ræturnar bólgna, missa rótarhárin og verða marg-greinóttar. Svepprótar-myndunin er mörgum sveppum nauðsynleg, því að þeir fá sykrur úr trjánum. Samlífi svepps og trés er þó flóknara en svo og má líta á það sem hag beggja, samhjálp, því að tréð fær vatn og nitursambönd frá sveppnum. Nærri öll skógartré lifa í sambýli við sveppi og sum tré við fleiri en eina tegund. – Einnig er til hin þriðja gerð af svepprót, innræn (endotrof), en þá eru ímur aðallega inni í sjálfum rótarfrumunum en lítið á milli á þeirra.

Jarðvegssveppir mynda oft svo kallaða sveppabauga (álfahringi). Sveppþalið gefur frá sér efni, sem smám saman dregur úr vextinum, svo að hann heldur aðeins áfram í hring umhverfis gamla þalið. Sveppurinn hefur líka áhrif á grasvöxt og örvast vöxturinn einkum þar, sem nitursambönd losna helst. Gamalt sveppþal veldur hins vegar því, að verulega dregur úr grasvexti. Orsök þess er ókunn. Þessu fylgir mismunur í grasvexti, svo að myndast hringir, nornabaugar, sem geta orðið nokkrir metrar að þvermáli.

 

Þverskurður af hálfum nornabaug. 1. eðlilegur vöxtur inn við miðju, 2. grösugur innri hringur, 3. gróðurvana belti, 4. grösugur ytri hringur, 5. eðlilegur vöxtur utan hrings, m. sveppímur í jarðvegi.

Þverskurður af hálfum nornabaug. 1. eðlilegur vöxtur inn við miðju, 2. grösugur innri hringur, 3. gróðurvana belti, 4. grösugur ytri hringur, 5. eðlilegur vöxtur utan hrings, m. sveppímur í jarðvegi.

1.8 Vanburðasveppir (Fungi imperfecti)

Allmargar tegundir sveppa mynda ýmist engin gró eða aðeins kóníðugró. Þeir eru sagðir ófullkomnir (»imperfect«) eða vanburða, því að kynhirzlur eru óþekktar, svo að ekki er unnt að flokka þá með öðrum tegundum, sem fjölgar með kynæxlun (eru »perfect«). Nú hefur verið lýst um 25’000 tegundum, þar sem kynæxlun er óþekkt. Einstaka sinnum hefur tekist að finna eina og eina tegund, þar sem kynæxlun kemur fyrir, og hefur tegundin þá verið flutt á sinn stað í kerfinu, en þó eru þær oft látnar óhreyfðar af handhægum ástæðum. Það hefur komið í ljós, að flestar tegundir vanburðarsveppa teljast til asksveppa en nokkrar þó til kólfsveppa eins og sjá má á gerð sveppþráða. Kynæxlunarlíki er allalgengt meðal vanburðasveppa og sýnist koma að nokkru leyti í stað kynæxlunar. Hópurinn vanburðasveppir er ekki fræðileg flokkunareining innan kerfisfræðinnar en samt má líta á hópinn sem flokk eða fylkingu.

 

 

1.9 Sveppir og menn

Enda þótt ýmsir sveppir séu mönnum til mikils miska og sumir hverjir lífshættulegir, eru aðrir til verulegs ávinnings. Allmargar tegundir eru til mikilla nytja, því að þær framleiða efni, sem víða koma að gagni. Þeir hafa verið notaðir frá ómuna tíð og enn er verið að uppgötva ný efni í þeim. Gersveppir eru notaðir við ostagerð og bakstur, meðal annars vegna þess að þeir mynda koldíoxíð (CO2), og þeir mynda líka etanól (vínanda). Ýmis efni í sveppum eru notuð sem lyf eins og penísillín, sem var fyrsta fúkalyfið.

Árið 1979 var hafin framleiðsla á lyfinu syklóspóríni, en það er einangrað úr jarðvegssveppi (Tolypocladium inflatum). Þetta er hringsameind, gerð úr 13 amínósýrum, og hefur ein sýran hvergi fundizt annars staðar í náttúrunni. Syklóspórínið er gætt þeim eiginleika að bæla ónæmiskerfi líkamans, svo að meiri líkur eru á því en áður að líkami sjúklings hafni ekki ígræddu líffæri. Önnur hliðstæð lyf hafa þann ókost að drepa mergfrumur í beini, svo að sjúklingar fengu oft hvítblæði í kjölfar ígræðslu.

Vanburðasveppir hafa mikla hagnýta þýðingu. Til að mynda eru fáeinar tegundir af ættkvíslinni Penicillium (pensilhý), sem gefa osti sérkennilegt bragð og ilm að dómi sælkera. Sú saga er sögð, að bóndasonur hafi gleymt nestispakka sínum í helli skammt frá þorpinu Roquefort í Frakklandi. Þegar hann náði í nestið að nokkrum vikum liðnum var osturinn í pakkanum orðinn ilmandi og morkinn. Þótti hann svo góður, að menn tóku að framleiða slíkan ost, sem nú gengur undir nafni dregið af heiti þorpsins og er öðrum óheimilt að nota það. Sveppurinn, sem breytingunum olli og þrífst þarna í hellum, heitir Penicillium roquefortii. Annar sveppur af sömu ættkvísl er P. camembertii, sem gefur kamemberosti sérlegt bragð. Í Austurlöndum fjær eru bursthýstegundir, Aspergillus oryzae og A. soyae, notaðar ásamt mjólkursýrugerlum til að búa til ýmsa sojabaunarétti. Tegundin A. oryzae er líka notuð við framleiðslu á hinu þekkta japanska hrísgrjónavíni ásamt Saccharomyces cerevisiae. Sítrónusýra er framleidd á þann hátt, að bursthý (Aspergillus) er látið vaxa í breiðum við mjög súrar aðstæður.

Fúkalyf eru efni sem myndast í einni lífveru en hefta vöxt annarrar (eins og gerla) og koma sum þeirra mönnum til góða. Allmörg fúkalyf eru framleidd úr vanburðasveppum. Fyrsta fúkalyfið var uppgötvað 1928. Þá vildi svo til, að sveppurinn Penicillium notatum blandaðist við Staphylococcus gerla, sem verið var að rækta á næringaræti. Vöxtur gerlanna stöðvaðist algerlega og fengu menn þá hugmynd um að nota sveppinn sem lyf. Tíu árum seinna tókst að framleiða hreint penísillín og hafin var stórfelld framleiðsla á því. Í síðari heimsstyrjöldinni jókst framleiðsla þess gífurlega enda reyndist það frábærlega vel við ýmsum sjúkdómum, sem gram-jákvæðir gerlar valda. Af þeim má nefna sárasótt (sýfilis), lekanda, lungnabólgu og barnaveiki.

Sum efni í sveppum eru mjög eitruð og geta líka valdið krabbameini, eins og aflatoxín, sem myndast í vissum stofnum af Aspergillus flavus og A. paraciticus. Sveppir þessir ná á stundum að vaxa í fæðuefnum (t.d. jarðhnetum) og valda miklu heilsutjóni.

Húðsveppir (dermatophytes; úr grísku dermatos, skinn og phyton, planta) eru oft taldir til sérstaks hóps innan vanburðasveppa. Þeir valda oft þrálátum sjúkdómum í húð, svo sem hringormi (reformi), geitum og fótsveppum. Einkum eru sjúkdómar þessir algengir í löndum hitabeltis. Flesta sveppina hefur tekizt að ákvarða sem asksveppi, en þar sem þeir valda sjúkdómum á kynlausu æviskeiði sínu, eru þeir enn taldir til vanburðasveppa. Hringormur lýsir sér sem hringmyndaðir, hreistursblettir með lítið eitt dökkleitari upphækkuðum garði í jöðrunum. Nokkrar tegundir, sem lifa á mönnum, þrífast líka á skepnum. Skemmst er að minnast sveppasýkingar af völdum Trichophyton mentagrophytes, sem varð vart í hrossum hér á landi sumarið 1992.

Sveppir þrífast bezt á rakri og heitri húð, til að mynda undir brjóstum og í leggöngum kvenna, undir höndum, í lærkrikum og á milli tánna. Sveppirnir gefa frá sér efni, sem drepa húðgerla, en þeir eru venjulega um 2 milljónir á hverjum fersentímetra og verja húðina. Þessir sveppir uxu líka á rökum trégólfum á ýmsum sundstöðum hér fyrr á árum og smituðust margir við að ganga þar berfættir. Víða var komið fyrir grunnum pönnum með 1% lausn af natríumhypoklóríði, sem menn voru látnir vaða yfir bæði fyrir og eftir sundspretti. Sveppirnir nærast á trefjaprótíninu keratíni í húðinni.

 

Þá má að lokum minnast á matsveppi. Neysla þeirra hefur mikið aukizt hin síðari ár. Íslenzka sveppaflóran hefur auðgazt mikið af gómsætum sveppum með aukinni ræktun trjáa á síðustu áratugum.

 

Kúalubbi (Leccinum scabrum) er algengur sveppur og góður á bragðið. Hann maðkar þó fljótt og stafur verður trefjakenndur. Ljósm. ÁHB.

Kúalubbi (Leccinum scabrum) er algengur sveppur og góður á bragðið. Hann maðkar þó fljótt og stafur verður trefjakenndur. Ljósm. ÁHB.

 

Berserkjasveppur (Amanita muscaria) er talinn eitraður. Hann vex í skógum, kjarri og hrísmóum. Hann vex víða um land en er algengastur á Norðausturlandi. Ljósm. ÁHB.

Berserkjasveppur (Amanita muscaria) er talinn eitraður. Hvítu hattflyksurnar eru leifar af heildarhulu. Hann vex í skógum, kjarri og hrísmóum. Hann vex víða um land en er algengastur á Norðausturlandi. Ljósm. ÁHB.

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