# Gljive

| Gljive Vremenski raspon: Rani devon, prije 410 miliona godina → Sadašnjost |
| --- |
| *Amanita muscaria*, *Sarcoscypha coccinea*, *Rhizopus stolonifer*, *Chytridiomycota*, *Penicillium*. |
| Sistematika |
| Domena | Eukaryota |
| Divizija |
| Blastocladiomycota
Chytridiomycota
Glomeromycota
Microsporidia
Neocallimastigomycota
Dikarya (inc. Deuteromycota)
Ascomycota
Pezizomycotina
Saccharomycotina
Taphrinomycotina
Basidiomycota
Agaricomycotina
Pucciniomycotina
Ustilaginomycotina
Potkoljena *incertae sedis*
Entomophthoromycotina
Kickxellomycotina
Mucoromycotina
Zoopagomycotina |

**Gljive** (latinski: *Fungi*) − označavane i kao **prave gljive** ili **Eumycota** − spadaju u najrasprostranjenije žive organizme na Zemlji. Smatra se da ih ima između 70.000 i 100.000 tzv. nižih i viših vrsta gljiva, mada najnovija istraživanja tu hipotetičku brojku pomjeraju na preko milion. Pretpostavlja se da svaku od pet stotina hiljada viših biljnih vrsta prati jedna do dvije vrste gljiva, pa računicu nije bilo teško izvesti.

Nauka o gljivama - mikologija (od grčkog myko-šampinjon, gljiva, logos-nauka) prošla je dug razvojni put. Njenim rodonačelnikom smatra se grčki filozof Aristotel, koji je dao prve opise gljiva.

Gljive su neobičan, raznovrstan i nedovoljno proučen svijet. Budući da ne vrše osnovnu funkciju biljaka fotosintezu jer ne posjeduju hlorofila, a ne kreću se i ne razmnožavaju polno poput životinja ne mogu se svrstati ni u prvo ni u drugo carstvo, već predstavljaju carstvo za sebe. One predstavljaju zaseban oblik života na zemlji koji je razvio tri osnovne strategije preživljavanja u prirodi a to su:

Gljive *Simbionti* koje žive u zajednici sa mnogim biljnim vrstama i međusobno se ispomažu. Gljive zadržavaju vodu u miceliju, a tu vodu koriste biljke koje pomoću korijenja izvlače mineralne materije neophodne za rast gljiva.

Gljive *Saprobionti* (razlagači) bez čijeg djelovanja u prirodi ne bi moglo doći do razlaganja organskih materija biljnog porijekla. i kaže se da nema gljiva šume bi se ugušile u vlastitom otpadu.

Gljive *Paraziti* koje napadaju žive biljke, životinje, ljude i druge gljive. Kroz mrežu tankih hranidbenih niti, gljive upijaju hranu iz žive ili nežive organske tvari.

Hifa je hranidbena nit gljive. Na vrhu hife nalaze se zaobljeni izdanci zvani haustoriji koji prodiru u ćelije, upijaju hranu i njome hrane gljivu.

Micelij je mreža hifa koja poput paukove mreže, prorasta supstrat i može se prostirati na više hektara površine.

Plodišta razvijaju gljive (makromicete) da bi se razmnožavale. Svako plodište oslobađa sićušne spore. Jedno plodište livadske pečurke proizvede oko 16 milijardi spora.

Klijanjem spora na odgovarajućoj podlozi nastaje novi micelij.

U odnosu na čovjeka, gljive mogu biti:

* *bezuslovno jestive* (upotrebljive u prijesnom stanju)
* *uslovno jestive* (upotrebljive samo ako su termički obrađene)
* *nejestive* (zbog ukusa ili konzistencije)
* *uslovno otrovne* (ako se konzumiraju s alkoholom)
* *bezuslovno otrovne i ljekovite*

## Evolucija

*Prototaxites milwaukeensis* (Penhallow, 1908),---
 srednjedevonska gljiva iz Wisconsina
Za razliku od evolucijska historije biljaka i životinja, rani fosilni zapisi o gljivama su oskudni. Faktori koji vjerovatno doprinose njihovoj nedovoljnoj zastupljenosti među fosilima uključuju prirodu gljiva plodnuh tijela, koja su mehka, mesnata i lahko razgradljiva tkiva, kao i mikroskopske dimenzije njihovih struktura, koje stoga nisu lahko uočljive. Gljivne fosile je teško razlikovati od onih od drugih mikroba, a najlakše ih je prepoznati ako nalikuju postojećim gljivama. Često se nalaze i uzimaju iz mineraliziranih biljaka ili životinja – domaćina; ti se uzorci obično proučavaju izradom preparata tankog presjeka koji se mogu pregledati pomoću svjetlosne ili transmisijske eletronske mikroskopije. Istraživači proučavaju fosilne kompresije otapanjem okolne matrice kiselinom, a zatim pomoću svjetlosti ili skenirajućom elektronskom mikroskopijom da bi se uočili detalji površine.
Najraniji fosili koji imaju obilježja gljiva karakteristični za doba Paleoproterozoika, prije nekih 2.400 miliona godina; ti višećelijski, bentoski organizmi imali nitaste strukture sposobne za anastomozu.

Druge studije (2009) procenjuju dolazak gljivnih organizama na oko 760–1.060 miliona godina prije sadašnjosti, a na osnovu upoređivanja brzine evolucije u usko povezanim grupama. Za veći dio ere paleozoik (prije 542–251 milion godina), čini se da su gljive bile vodene i sastojale se od organizama sličnih postojećim Chyrididama koji imaju spore s bičem. Evolucijsko prilagođavanje, od vodenog na kopneni način života zahtijevalo je diverzifikaciju ekoloških strategija za dobijanje hranjivih sastojaka, uključujući parazitizam, saprobnost, te razvoj uzajamnosti, poput mikoriza i lihenizacija. Nedavna (2009) istraživanja sugeriraju da je ekološki status predaka Ascomycota bio je saprobizam, i neovisni događaji sa lišajima su se dogodili više puta.

U maju 2019., naučnici su izvijestili o otkriću gljive fosila nazvane *Ourasphaira giraldae* u kanadskom Arktiku, koja je na mogla rasti i prije milijardu godina, mnogo prije nego što su e biljke pojavile na kopnu.

## Taksonomija

Iako su obično uključene u nastavne programe i udžbenike o botanici, gljive su bliže životinjama nego biljkama i smještaju se sa životinjama u monofletnu grupu Opisthokonta. Analize koje koriste molekulsku filogenetiku podržavaju monofiletsko porijeklo gljiva. Taksonomija gljiva je u stanju stalnog mijenjanja, posebno zbog nedavnih istraživanja temeljenih na usporedbama sekvenci DNK. Nove filogenetske analize često poništavaju klasifikacije zasnovane na starijim, a ponekad i manje diskriminativnim metodima, utemeljenim na morfološkim obilježjima i konceptima bioloških vrsta dobivenih eksperimentalnim ukrštanjem.
Ne postoji jedinstvenći općenito prihvaćen sistem na razini višiih taksonomskih rangova i česte su promjene imena na svim razinama, od vrsta pa nadalje. U toku su napori među istraživačima da se uspostavi i ohrabri upotreba jedinstvene i konzistentnije nomenkature. Vrste gljiv mogu imati i više naučnih imena, ovisno o njihovom životnom ciklusu i načinu (seksualne ili aseksualne) reprodukcije. Web stranice poput Index Fungorum i ITIS (*Integrated Taxonomic Information System*) navode današnja imena vrsta gljiva (s unakrsnim referencama na starije sinonime).

Klasifikacija carstva Fungi iz 2007., rezultat je opsežnog istraživačkog napora koji uključuje desetine mikologa i drugih naučnika koji rade na taksonomiji gljiva. Prepoznaje sedam koljena, od kojih su dva – Ascomycota i Basidiomycota – u ogranku koji predstavlja potcarstvo Dikarya, najbogatiju i najpoznatiju grupu, uključujući sve gljive, najviše plijesni hrane, najviše patogenih gljiva biljaka i piva, vinoa i hljebni kvasac. Prateći kladogram prikazuje glavne taksone gljiva i njihov odnos prema opistokontnim i unikontnim organizmima, utemeljen na radu Philippea Silara, "Mycota: Sveobuhvatni traktat o gljivama kao eksperimentalnim sistemima za osnovna i primijenjena istraživanja" (*The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research*) i Tedersooa et al. 2018. Dužine grana nisu proporcionalne evolucijskim udaljenostima.

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| Zoosporia |

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| Rozellomyceta |

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| Rozellomycota |
Rozellomycetes |
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| Microsporidiomycota |

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|   | *Mitosporidium* |
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|   | *Paramicrosporidium* |
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|   | *Nucleophaga* |
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Metchnikovellea |
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Microsporea |
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| Aphelidiomyceta |

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| Aphelidiomycota |
Aphelidiomycetes |
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| Eumycota |

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| Chytridiomyceta |

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| Neocallimastigomycota |
Neocallimastigomycetes |
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| Chytridiomycota |

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| Monoblepharomycotina |

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Hyaloraphidiomycetes |
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Monoblepharidomycetes |
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Sanchytriomycetes |
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| Chytridiomycotina |

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Mesochytriomycetes |
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Chytridiomycetes |
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| Blastocladiomyceta |

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| Blastocladiomycota |

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Blastocladiomycetes |
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Physodermatomycetes |
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| Amastigomycota |

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| Zoopagomyceta |

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| Basidiobolomycota |

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Basidiobolomycetes |
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Olpidiomycetes |
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| Entomophthoromycota |

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Neozygitomycetes |
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Entomophthoromycetes |
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| Kickxellomycota |

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| Zoopagomycotina |
Zoopagomycetes |
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| Kickxellomycotina |

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Dimargaritomycetes |
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Kickxellomycetes |
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| Mortierellomycota |
Mortierellomycetes |
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| Mucoromyceta |

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| Calcarisporiellomycota |
Calcarisporiellomycetes |
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| Mucoromycota |

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Umbelopsidomycetes |
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Mucoromycetes |
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| Symbiomycota |

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| Glomeromycota |

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Paraglomeromycetes |
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Archaeosporomycetes |
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Glomeromycetes |
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| Dikarya |

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| Entorrhizomycota |
Entorrhizomycetes |
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Basidiomycota |
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Ascomycota |
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| Basidiomycota |

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| Pucciniomycotina |

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Tritirachiomycetes |
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Mixiomycetes |
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Agaricostilbomycetes |
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Cystobasidiomycetes |
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Classiculaceae |
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Microbotryomycetes |
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Cryptomycocolacomycetes |
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Atractiellomycetes |
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Pucciniomycetes |
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| Orthomycotina |

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| Ustilaginomycotina |

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Monilielliomycetes |
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Malasseziomycetes |
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Ustilaginomycetes |
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Exobasidiomycetes |
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| Agaricomycotina |

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| --- | --- |
|   |
?Geminibasidiomycetes |
|   |
|   |
?Wallemiomycetes |
|   |
|   |
Bartheletiomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Tremellomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Dacrymycetes |
|   |
|   |
Agaricomycetes |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Ascomycota |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Taphrinomycotina |

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |
| --- | --- |
|   |
Neolectomycetes |
|   |
|   |
Taphrinomycetes |
|   |

 |
|   |
| Schizosaccharomyceta |

|  |  |
| --- | --- |
|   |
Archaeorhizomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Pneumocystidomycetes |
|   |
|   |
Schizosaccharomycetes |
|   |

 |
|   |

 |
|   |

 |
|   |
| Saccharomyceta |

|  |  |
| --- | --- |
| Saccharomycotina |
Saccharomycetes |
|   |
| Pezizomycotina |

|  |  |
| --- | --- |
|   |
?Thelocarpales |
|   |
|   |
?Vezdaeales |
|   |
|   |
?Lahmiales |
|   |
|   |
?Triblidiales |
|   |
|   |
Orbiliomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Pezizomycetes |
|   |
| Leotiomyceta |

|  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- | --- |
| Sordariomyceta |

|  |  |
| --- | --- |
|   |
Xylonomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Geoglossomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Leotiomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Laboulbeniomycetes |
|   |
|   |
Sordariomycetes |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |
| Dothideomyceta |

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |
| --- | --- |
|   |
Coniocybomycetes |
|   |
|   |
Lichinomycetes |
|   |

 |
|   |
|   |

|  |  |  |  |  |  |  |  |
| --- | --- | --- | --- | --- | --- | --- | --- |
|   |

|  |  |
| --- | --- |
|   |
Eurotiomycetes |
|   |
|   |
Lecanoromycetes |
|   |

 |
|   |
|   |

|  |  |
| --- | --- |
|   |
Collemopsidiomycetes |
|   |
|   |

|  |  |
| --- | --- |
|   |
Arthoniomycetes |
|   |
|   |
Dothideomycetes |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

 |
|   |

## Klasifikacija

Klasifikacija gljiva je prošla dug razvojni put. U početku je u grupu gljiva uključivano mnogo grupa koje se danas svrstavaju u druga carstva. Najsavremenija istraživanja dovode u blisku filogenetsku vezu gljive i životinje te ih uključuje u zajedničku grupu pod nazivom Opisthokonta.

## Ljudska upotreba

Ćelije *Saccharomyces cerevisiae* prikazane uz pomoć DIC mikroskopije
Ljudska upotreba gljiva za pripremu ili konzerviranje hrane i druge svrhe je opsežna i ima dugu historiju. Uzgoj gljiva i sakupljanje gljiva su velike industrije u mnogim zemljama. Izučavanje historijskih upotreba i sociološkog utjecaja gljiva poznata je kao etnomikologija. Zbog sposobnosti ove grupe da proizvede ogroman asortiman prirodnih proizvoda s antimikrobnim ili drugim biološkim djelovanjima, mnoge vrste se već dugo koriste ili se razvijaju za industrijsku proizvodnju antibiotika, vitamina i lijekova protiv raka i za smanjenje holesterola. Razvijene su metode za genetičko inženjerstvo gljiva, omogućavajući metaboličko inženjerstvo gljivičnih vrsta. Na primjer, genetska modifikacija vrsta kvasca, koje je lako uzgajati velikom brzinom u velikim posudama za fermentaciju, otvorila je načine farmaceutske proizvodnje koji su potencijalno efikasniji od proizvodnje izvornih organizama. Industrije zasnovane na gljivama ponekad se smatraju glavnim dijelom rastuće bioekonomije, s primjenama u fazi istraživanja i razvoja, uključujući upotrebu za tekstil, zamjenu mesa i opću biotehnologiju gljiva.

## Reference

## Vanjski linkovi

*  Arhivirano 25. 1. 2021. na Wayback Machine
*  Arhivirano 2. 5. 2020. na Wayback Machine
* Luketa Stefan: Carstvo gljiva Arhivirano 11. 11. 2009. na Wayback Machine, BioNet škola, 2009.
*  Prvi web site o gljivama u BiH (bs)
* The WWW Virtual Library: Mycology (en)
* MykoWeb (en)
* Morchella Fungi Site Arhivirano 23. 2. 2009. na Wayback Machine (bs)