06.04.2013 Views

Fungi – moulds, yeasts and mushrooms

Fungi – moulds, yeasts and mushrooms

Fungi – moulds, yeasts and mushrooms

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

<strong>Fungi</strong> <strong>–</strong><br />

<strong>moulds</strong>, <strong>yeasts</strong> <strong>and</strong> <strong>mushrooms</strong><br />

© Anna Gorbushina, WS 2006/2007<br />

<strong>Fungi</strong> in microbiology<br />

(probably, you have heard these names already…)<br />

Applied microbiology <strong>and</strong> biotechnology<br />

Saccharomyces cerevisiae (beer, bread)<br />

Penicillium roquefortii (cheese)<br />

Penicillium chrysogenum <strong>and</strong> Acremonium chrysogenum<br />

(antibiotics)<br />

Aspergillus niger (citric acid)<br />

Haploid eukaryotes with fast growth rates<br />

Neurospora crassa<br />

Aspergillus nidulans<br />

Saccharomyces cerevisiae<br />

Ecology, physiology, diversity, plant pathology etc.<br />

© Anna Gorbushina, WS 2006/2007<br />

1


•Prokaryotes are morphologicall simple, but physiologically tremendously diverse<br />

•Planta, <strong>Fungi</strong> <strong>and</strong> Animalia have lived through this 3 domain transfromation with<br />

only minor alterations<br />

•Protists stay the most diverse group<br />

•<strong>Fungi</strong> traditionally includ true fungi <strong>and</strong> fungi-like protists<br />

Bacteria<br />

Archaea<br />

unicellular osmotrophic<br />

Protista unicellular diverse:<br />

ingestion, osmotrophic<br />

Animalia multicellular phagotrophic (ingestion)<br />

<strong>Fungi</strong> X-cellular osmotrophic<br />

Planta multicellular osmotrophic<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

2


<strong>Fungi</strong> - what is treated by<br />

mycology?<br />

True fungi <strong>and</strong> fungi-like protists<br />

Microscopic organisms, but there are also<br />

huge fungi<br />

For classification it is necessary to have<br />

propagation structures or to use molecular<br />

biological characters<br />

© Anna Gorbushina, WS 2006/2007<br />

<strong>Fungi</strong> <strong>and</strong> mycology<br />

Groups traditionally treated by mycology<br />

Oomycetes<br />

Chytridiomycetes<br />

Zygomycetes<br />

Ascomycetes<br />

Basidiomycetes<br />

© Anna Gorbushina, WS 2006/2007<br />

3


Protozoa<br />

Mycetozoa<br />

Mesomycetozoa<br />

Chromista<br />

Oomycota<br />

Eumycota<br />

New classification<br />

Kingdom Protista<br />

Plasmodiophoromycota<br />

Dictyosteliomycota<br />

Acrasiomycota<br />

Myxomycota<br />

Oomycota<br />

Hyphochytriomycota<br />

Labyrinthulomycota<br />

Chytridiomycota<br />

Zygomycota<br />

Zygomycetes<br />

Entomophthorales<br />

Mortierellales<br />

Mucorales<br />

Ascomycota<br />

Archiascomycetes<br />

Pneumocystidales<br />

Hemiascomycetes<br />

Saccharomycetales<br />

Euascomycetes<br />

Chaetothyriales<br />

Clavicipitales<br />

Dothideales<br />

Eurotiales<br />

Hypocreales<br />

Leotiales<br />

Microascales<br />

Onygenales<br />

Ophiostomatales<br />

Phyllachorales<br />

Pleosporales<br />

Sordariales<br />

Basidiomycota<br />

Hymenomycetes<br />

Agaricales<br />

Stereales<br />

Tremellales<br />

Urediniomycetes<br />

Sporidiales<br />

Ustilaginomycetes<br />

Microstomatales<br />

Tilletiales © Anna Gorbushina, WS 2006/2007<br />

Ustilaginales<br />

Kingdom <strong>Fungi</strong><br />

Chytridiomycota<br />

Zygomycota<br />

Ascomycota<br />

Basidiomycota<br />

© Anna Gorbushina, WS 2006/2007<br />

4


© Anna Gorbushina, WS 2006/2007<br />

<strong>Fungi</strong> or How to define a diverse polyphyletic<br />

group (i)<br />

Heterotrophic (no photosynthesis)<br />

Osmotrophic (absorptive nutrition -<br />

exoenzymes)<br />

Cell walls with chitin<br />

Thallus grows in or on a substrate, is<br />

filamentous (or unicellular) <strong>and</strong> grows in<br />

modules<br />

Eukaryotic<br />

real nucleus, cells frequently multinuclear<br />

© Anna Gorbushina, WS 2006/2007<br />

5


<strong>Fungi</strong> or How to define a diverse polyphyletic<br />

group (ii)<br />

Life cycles diverse (from simple to<br />

complex)<br />

Sexual propagation may be present or<br />

absent<br />

Fruiting bodies (sporocarps) micro- or<br />

macroscopic, sometimes differentiated into<br />

tissue-like structures<br />

<strong>Fungi</strong><br />

eukaryotic, haploid (n)<br />

dikaryotic (n+n) <strong>and</strong> diploid (2n)<br />

periods<br />

unmotile<br />

an exception in <strong>Fungi</strong>: spores of<br />

Chytridiomycota<br />

Cell walls predominantly contain chitin<br />

(R=NH-CO-CH 3 )<br />

unicellular (yeast) <strong>and</strong> multicellualr<br />

(filamentous or mycelial fungi)<br />

Apical (tip) growth<br />

Spores as spreading <strong>and</strong> survival units<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

6


Growth in hypha <strong>and</strong> yeast<br />

Fungal spores<br />

From Bartnicki-Garcia & Lippman, 1969, © AAAS<br />

Spread (propagation) <strong>and</strong> survival<br />

spread - quantity<br />

Survival - thick pigmented cell walls<br />

Asexual (mitotic division)<br />

Sexual (cell fusion <strong>and</strong> meiosis)<br />

Sometimes in fruit bodies<br />

Protection<br />

Spreading mechanisms<br />

Apical growth<br />

Transport<br />

system to the<br />

only growth<br />

zone - the tip<br />

Numerous<br />

vesicles with<br />

„building units“<br />

for cell wall<br />

synthesis<br />

Spitzenkörper<br />

Yeast - different<br />

distribution of<br />

growth zones<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

7


Fungal fruiting bodies (sporocarps)<br />

Mushrooms-onlyfruit<br />

bodies<br />

Still composed of<br />

hyphae<br />

From Cooke&Rayner, 1984 (c) Longman<br />

© Prof. Trinci<br />

© Anna Gorbushina, WS 2006/2007<br />

Morphology of fungi - hypha, mycelium, colony<br />

•Mycelium has 1 to 10 µm in diameter<br />

•Tip growth<br />

•Substrate penetration<br />

•Substrate contact surface is enormous<br />

From Cooke&Rayner, 1984 (c) Longman<br />

© Polona Zalar & Nina Gunde-Cimerman<br />

© Anna Gorbushina, WS 2006/2007<br />

8


Morphology of fungi - vegetative mycelium<br />

modifications<br />

Fungal morphology<br />

Witch circle<br />

Mycelial str<strong>and</strong>s -<br />

Serpula lacrymans<br />

Sclerotia - mummified fruits,<br />

insect bodies, with mycelium<br />

penetrating all through<br />

© Anna Gorbushina, WS 2006/2007<br />

© Polona Zalar & Nina Gunde-Cimerman<br />

© Anna Gorbushina, WS 2006/2007<br />

9


General information<br />

Cell wall - chitin, cellulose, glucanes<br />

Motility: none (flagellate spores in Chytridiomycota!)<br />

Thallus - mycelial (septate, unseptate) or yeast<br />

Metabolism: aerobic, facultative anaeroby<br />

(fermentation)<br />

Life cycles<br />

Asexual (vegetative, mitotic spores)<br />

Sexual (generative, meiotic spores)<br />

Complex, usually facultative<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

10


Oomycota<br />

Protozoa<br />

Mycetozoa<br />

Mesomycetozoa<br />

Chromista<br />

Oomycota<br />

Eumycota<br />

Chytridiomycota<br />

Zygomycota<br />

Zygomycetes<br />

Entomophthorales<br />

Mortierellales<br />

Mucorales<br />

Ascomycota<br />

Archiascomycetes<br />

Pneumocystidales<br />

Hemiascomycetes<br />

Saccharomycetales<br />

Euascomycetes<br />

Chaetothyriales<br />

Clavicipitales<br />

Dothideales<br />

Eurotiales<br />

Hypocreales<br />

Leotiales<br />

Microascales<br />

Onygenales<br />

Ophiostomatales<br />

Phyllachorales<br />

Pleosporales<br />

Sordariales<br />

Basidiomycota<br />

Hymenomycetes<br />

Agaricales<br />

Stereales<br />

Tremellales<br />

Urediniomycetes<br />

Sporidiales<br />

Ustilaginomycetes<br />

Microstomatales<br />

Tilletiales © Anna Gorbushina, WS 2006/2007<br />

Ustilaginales<br />

•Aquatic organisms<br />

•Water absolutely<br />

necessary (in soil, as<br />

l<strong>and</strong> plants parasites)<br />

•Heterokont, biflagellate,<br />

spores<br />

•Cell walls contain<br />

cellulose und glucan (ß-<br />

(1-3) <strong>and</strong> ß-(1-6)<br />

glycoside bonds<br />

•Oogamy with a complex<br />

<strong>and</strong> very specific<br />

fertilisation process<br />

•Phytophtora infestans<br />

© Anna Gorbushina, WS 2006/2007<br />

11


Protozoa<br />

Mycetozoa<br />

Mesomycetozoa<br />

Chromista<br />

Oomycota<br />

Eumycota<br />

Chytridiomycota<br />

Zygomycota<br />

Zygomycetes<br />

Entomophthorales<br />

Mortierellales<br />

Mucorales<br />

Ascomycota<br />

Archiascomycetes<br />

Pneumocystidales<br />

Hemiascomycetes<br />

Saccharomycetales<br />

Euascomycetes<br />

Chaetothyriales<br />

Clavicipitales<br />

Dothideales<br />

Eurotiales<br />

Hypocreales<br />

Leotiales<br />

Microascales<br />

Onygenales<br />

Ophiostomatales<br />

Phyllachorales<br />

Pleosporales<br />

Sordariales<br />

Basidiomycota<br />

Hymenomycetes<br />

Agaricales<br />

Stereales<br />

Tremellales<br />

Urediniomycetes<br />

Sporidiales<br />

Ustilaginomycetes<br />

Microstomatales<br />

Tilletiales © Anna Gorbushina, WS 2006/2007<br />

Ustilaginales<br />

Differences in<br />

propagation<br />

Sexual<br />

propagation (all<br />

around meiosis)<br />

Generations<br />

(sexual/asexual)<br />

© Anna Gorbushina, WS 2006/2007<br />

12


Some groups of <strong>Fungi</strong><br />

Chytridiomycota<br />

Zygomycota<br />

Ascomycota<br />

Basidiomycota<br />

Deuteromycota (<strong>Fungi</strong><br />

imperfecti)<br />

© Anna Gorbushina, WS 2006/2007<br />

Cell wall composition in phyla of fungi <strong>and</strong> fungallike<br />

protists<br />

Oomycota<br />

Chytridiomycota<br />

Zygomycota<br />

Ascomycota<br />

Basidiomycota<br />

cellulose<br />

β-(1-3)- <strong>and</strong> β-(1-6)glucan<br />

chitin<br />

β-(1-3)- <strong>and</strong> β-(1-6)glucan<br />

chitin<br />

chitosan<br />

chitin<br />

β-(1-3)- <strong>and</strong> β-(1-6)glucan<br />

chitin<br />

β-(1-3)- <strong>and</strong> β-(1-6)glucan<br />

glucan<br />

glucan<br />

polyglucouronate,<br />

glucuronomannproteins,<br />

polyphosphates<br />

galactomannoproteins<br />

α-(1-3)- glucan<br />

xylomannoproteins,<br />

α-(1-3)- glucan<br />

© Anna Gorbushina, WS 2006/2007<br />

13


Fungal nutrition <strong>and</strong> their<br />

presence in ecosystems<br />

Osmotrophic - with a huge reactive surface<br />

Organoheterotrophic<br />

Respiration/Fermentation<br />

Where they live: everywhere (soil, air, water,<br />

wood, rocks). Mainly aerated zones<br />

Strictly anaerobic fungus: only one<br />

Chytridiomycete Neocallimastix frontalis (39°C, pH<br />

6,5) known from the rumenal ecosystem<br />

© Anna Gorbushina, WS 2006/2007<br />

Hypha <strong>–</strong> mycelum <strong>–</strong> colony <strong>–</strong> fruiting body<br />

Mycelium important for all: parasites,<br />

symbionts, destruents<br />

Penetration <strong>and</strong> substrate colonisation<br />

Curious example: Nematode-trapping<br />

fungi<br />

© Anna Gorbushina, WS 2006/2007<br />

14


Fungal lifestyles<br />

Saprophytes on dead organic matter - „trash<br />

<strong>and</strong> litter“ :<br />

soil (1g of agricaltural field soil contains<br />

10-100 m of fungal mycelium)<br />

wood (destruction <strong>and</strong> staining)<br />

various organic compounds<br />

Symbionts with bacteria, algae, plants<br />

Parasites on animals, algae, plants<br />

SAPROPHYTES<br />

•Substrate mycelium, exoenzymes,<br />

osmotrophy<br />

•Available nutrients<br />

© Anna Gorbushina, WS 2006/2007<br />

-carbohydrates including highly polymeric<br />

(cellulose, starch, pectine, lignine)<br />

-proteins (including keratin)<br />

-fats, polyols etc<br />

© Anna Gorbushina, WS 2006/2007<br />

15


PARASITES<br />

•Direct contact <strong>and</strong> nutritional<br />

connection<br />

•Mycelium on the surface of a<br />

structure (e.g. leaf) or between the<br />

cells<br />

•Always a direct contact between<br />

cytoplasmatic membranes -<br />

penetration structure „haustorium“<br />

PARASITES<br />

•Metabiotrophic<br />

•Biotrophic<br />

•Necrotrophic<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

16


Claviceps purpurea - sclerotia<br />

SYMBIONTS<br />

Obligate life cycle -<br />

perfectly adapted to the<br />

host‘s life cycle<br />

Infection is very limited -<br />

only one corn<br />

Sclerotia are survival<br />

structures (winter winter), ), after<br />

that a proper meiotic<br />

cycle takes place<br />

Various biologically active<br />

metabolites (LSD)<br />

Mycorrhiza <strong>–</strong> higher plants (trees, grasses,<br />

orchids)<br />

Lichens <strong>–</strong> algae <strong>and</strong> /or cyanobacteria<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

17


Mycotrophy of higher plants<br />

• 80% of plant species are obligately mycotrophic, 10% -<br />

facultatively<br />

•<strong>Fungi</strong> form a close contact with plant roots either as a coat<br />

or between the root parenchyme cells<br />

•Characteristic morphology <strong>and</strong> physiology<br />

Mycorrhiza<br />

Mutual profit<br />

Water, P-, N- <strong>and</strong> Ca-compounds for a<br />

plant partner<br />

Carbohydrate for a fungal partner<br />

What happens on a contact interface?<br />

Hyphae are digested<br />

Phosphatases of plant<br />

Glucose und succrose are transformed into<br />

trehalose und mannitol<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

18


Ectotrophic mycorrhiza<br />

Fs <strong>–</strong> fungal sheath (Mantel aus Pilzhyphen, die auch<br />

außerhalb der Wurzel weiter in den Boden wachsen)<br />

Hn <strong>–</strong> Hartig‘s net (Myzel zwischen den Zellen)<br />

Mycorrhiza stimulates growth of plants<br />

© Anna Gorbushina, WS 2006/2007<br />

See also: Smith, S.E., Smith, F.A. & Jakobsen, I. (2003) Plant Physiol. 133: 16-20.<br />

© Anna Gorbushina, WS 2006/2007<br />

19


Ectotrophic<br />

mycorrhiza makes<br />

phosphorous<br />

available, also<br />

including<br />

phosphates from<br />

mineral soil<br />

horizons<br />

L<strong>and</strong>eweert, R., Hoffl<strong>and</strong>, E., Finlay, R.D., Kuyper, T.W. & Van Breemen, N.<br />

(2001) Trends Ecol. Evol. 248-253.<br />

© Anna Gorbushina, WS 2006/2007<br />

Lichen thallus<br />

Form <strong>and</strong> structure are<br />

determined by a mycobiont<br />

Cortex (upper <strong>and</strong> lower)<br />

Photobiont layer<br />

Medula<br />

Flavoparmelia caperata<br />

© Anna Gorbushina, WS 2006/2007<br />

20


<strong>Fungi</strong><br />

Diverse<br />

Ubiquitous <strong>and</strong> stress-tolerant<br />

With a huge potential (enzymes,<br />

secondary metabolic products)<br />

Important in nature as destruents<br />

(recyclers), symbionts <strong>and</strong> parasites; as<br />

lichens even as primary producers<br />

Biotechnologically useful<br />

© Anna Gorbushina, WS 2006/2007<br />

In the real world, the estimate of fungus biomass<br />

is about two tons per human being in the overall<br />

biosphere. The unseen underground feeding<br />

mechanisms of fungi are the great recyclers:<br />

autumn leaves, cornstalk debris, mountains of<br />

trash, all returned to usable form mainly by fungi.<br />

No soil fungi, no trees <strong>and</strong> forests as we know<br />

them. No waving fields of grain. No escape from<br />

unforeseen pneumonia. No athlete's foot. No<br />

bread, no booze. Give us non-mushroom<br />

mycologists a break!<br />

Dr. Emory Simmons<br />

© Anna Gorbushina, WS 2006/2007<br />

21


Zygomycota<br />

Zygomycota<br />

Chitinous cell<br />

walls<br />

Spores in<br />

sporangia<br />

Survival<br />

structures -<br />

zygosporangia<br />

© Anna Gorbushina, WS 2006/2007<br />

Sugar fungi<br />

Insect<br />

parasites<br />

Symbionts<br />

(VAM<br />

Mykorrhiza)<br />

© Anna Gorbushina, WS 2006/2007<br />

22


Ascomycota<br />

The biggest group<br />

Chitin, all protective<br />

pigments<br />

Spores asexual <strong>–</strong><br />

exogenous; sexual <strong>–</strong> in<br />

Asci<br />

Ascomycota<br />

Parasites (true mildew)<br />

Symbionts (lichens,<br />

mykorrhiza)<br />

Destruents<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

23


Basidiomycota<br />

Chitin<br />

Dikaryotic<br />

phase<br />

(reproductio<br />

n process is<br />

prolonged)<br />

Spores<br />

exogenous<br />

on a<br />

basidium;<br />

basidia<br />

(plural) on a<br />

fruit body<br />

Basidiomycota<br />

Saprotrophs<br />

(wood<br />

decomposers)<br />

Parasites<br />

(living hosts,<br />

also trees)<br />

Symbiontes<br />

(mycorrhiza)<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

24


Protozoa<br />

Mycetozoa<br />

Mesomycetozoa<br />

Chromista<br />

Oomycota<br />

Eumycota<br />

Chytridiomycota<br />

Zygomycota<br />

Zygomycetes<br />

Entomophthorales<br />

Mortierellales<br />

Mucorales<br />

Ascomycota<br />

Archiascomycetes<br />

Pneumocystidales<br />

Hemiascomycetes<br />

Saccharomycetales<br />

Euascomycetes<br />

Chaetothyriales<br />

Clavicipitales<br />

Dothideales<br />

Eurotiales<br />

Hypocreales<br />

Leotiales<br />

Microascales<br />

Onygenales<br />

Ophiostomatales<br />

Phyllachorales<br />

Pleosporales<br />

Sordariales<br />

Basidiomycota<br />

Hymenomycetes<br />

Agaricales<br />

Stereales<br />

Tremellales<br />

Urediniomycetes<br />

Sporidiales<br />

Ustilaginomycetes<br />

Microstomatales<br />

Tilletiales © Anna Gorbushina, WS 2006/2007<br />

Ustilaginales<br />

Ameboid slime <strong>moulds</strong> - Protista<br />

neither slimy, no <strong>moulds</strong><br />

no hyphae<br />

no cell walls in the trophic phase<br />

in the trophic phase ingestion or<br />

phagocytosis<br />

fruiting bodies with cell walls<br />

Spores as propagation units<br />

amoebal or plasmodial vegetative phase<br />

© Anna Gorbushina, WS 2006/2007<br />

25


Ameboid slime <strong>moulds</strong> - Protista<br />

Acrasiomycota: early separation from<br />

Eukarya - discoidal mitochondrial cristae<br />

Myxomycota <strong>and</strong> Dictyosteliomycota <strong>–</strong><br />

later development, unrelated<br />

tubular mitochondrial crystae<br />

Numerous differences in morphology, life<br />

cycles, flagella, cell wall composition,<br />

nutrition<br />

© Anna Gorbushina, WS 2006/2007<br />

Plasmodium Physarum polycephalum<br />

© Anna Gorbushina, WS 2006/2007<br />

26


Different types of sporangia in<br />

myxomycetes<br />

Trichia decipiens<br />

Physarum cinereum<br />

Ameboid slime <strong>moulds</strong><br />

Pigments, heterotrophic (no<br />

chlorophyll)<br />

Net-work like structures<br />

that creep over substrates<br />

<strong>and</strong> are capable of<br />

phagocytosis<br />

rhythmic plasmamovements<br />

over cmdistances<br />

Fruiting bodies (sporangia)<br />

Diachea leucopodium<br />

Stemonitis<br />

fusca<br />

© Anna Gorbushina, WS 2006/2007<br />

© Anna Gorbushina, WS 2006/2007<br />

27

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!