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1 Zvyšování konkurenceschopnosti studentů oboru botanika a učitelství biologie CZ.1.07/2.2.00/ B. Mieslerová, A. Lebeda, (KB PřF UP v Olomouci)

2 GENERAL MYCOLOGY Prof. Ing. Aleš Lebeda, DrSc. RNDr. Barbora Mieslerová, Ph.D.

3 LIST OF RECOMMENDED LITERATURE Alexopoulos, C.J., Mims, C.W., Blackwell, M.: Introductory Mycology. John Wiley & Sons, New York, Deacon, J.W.: Modern Mycology. Blackwell Science Ltd., Oxford, lbiology/

4 Dix, N.J., Webster, J.: Fungal Ecology. Chapman & Hall, London, Carlile, M.J., Watkinson, S.C., Gooday, G.W.: The fungi. Academic Press, San Diego, Gryndler, M. (eds.): Mykorhizní symbióza. Academia, Griffin, D.H.: Fungal Physiology. Willey-Liss, New York, 1994.

5 Kalina, T., Váňa, J.: Sinice, řasy, houby, mechorosty a podobné organismy v současné biologii. Praha, Karolinum 2005, Isaac, S.: Fungal- Plant Interactions, Kluwer Academic Publishers, Kavanagh, K. (ed.): Fungi. Biology and Applications. John Wiley & Sons, 2005.

6 Ingold, C.T., Hudson, H.J. (1996): The biology of fungi. Chapman & Hall, London, UK. Klán, J.: Co víme o houbách. SPN Praha, 1989 Váňa, J.: Systém a vývoj hub a houbových organismů. UK, Praha, Kendrick, B The Fifth Kingdom, 2nd edition. Mycologue Publications, Waterloo, Ontario, Canada.

7 JOURNALS In the Czech Republic Česká mykologie Czech Mycology ČVSM Mykologické listy ČVSM Mykologický sborník ČMS

8 JOURNALS MYCOTAXON Order dept. Mycotaxon, USA MYCOLOGICAL RESEARCH Elsevier Science The Netherlands MYCOLOGIA APT Press

9 MYCOLOGIST Elsevier Science The Netherlands FIELD MYCOLOGY Elsevier Science The Netherlands FUNGAL GENETICS AND BIOLOGY Elsevier

10 FUNGAL CHARACTERIZATION Nutrition: Heterotrophic (photosynthesis lacking) and absorptive (ingestion rare), exceptionally fagocytosis. Thallus: Plasmodial, amoeboid or pseudoplasmodial Unicellular or philamentous (mycelial, septate or nonseptate) Nonmotile but motile states (zoosporas) may occur Cell wall: Well-defined, typicaly chitinised (cellulose in Oomycota), periplast (Myxomycota). Nuclear status: Eukaryotic, multinucleate, the mycelium being homo- or heterokaryotic, haploid, dikaryotic, or diploid, the last being usually of limited duration. Life cycle: Simple to complex. Sexuality: Asexual or sexual and homo- or heterothallic. Sporocarps: Microscopic or macroscopic and showing limited tissue differentiation. Habitat: Ubiquitous as saprofytes, symbionts, parasites or hyperparasites Distribution: Cosmopolitan

11 CLASSIFICATION OF ORGANISMS Whittaker s scheme (1969)

12 Classifications of all cellular organisms into three domains EUKARYA, BACTERIA and ARCHAEA (Cavalier- Smith, 1998).

13 Suffixes used to denote taxonomic ranks of fungal taxa Rank suffix Division (Phylum) Subdivision (Subphylum) Class Subclass Order Family - mycota - mycotina - mycetes - mycetidae - ales - aceae

14 SYSTEM OF FUNGI Corliss, 1994; Cavalier-Smith 1995 Kingdom: PROTOZOA Amoeboid or plasmodial organisms naked in somatic stage, flagella without mastigonemata, able to ingest particulate food Phylum: MYXOMYCOTA Amoeboid organisms or grow as a network of plasmodium, that engulfs bacteria, and then form fruiting bodies that release spores Phylum : PLASMODIOPHOROMYCOTA Obligate intracellular parasites of plantae, algae and fungi, exist as naked plasmodia in the host cells; form highly persistent thick-walled resting spores. Fuligo septica

15 Kingdom: CHROMISTA (STRAMENOPILA) One cell or hyphae organisms with cellulose-based walls and biochemical features resembling those of plants. Flagella with mastigonemata, nutrition exclusively absorptive Phylum : LABYRINTHULOMYCOTA Phylum : OOMYCOTA Hyphae aseptate; asexual reproduction by formation of motile, biflagellate zoospores in a sporangium; sexual reproduction by fusion of male sex organ (antheridium) and female sex organ (oogonium), leading to production of thick-walled resting spores (oospores) Phylum : HYPHOCHYTRIOMYCOTA Pseudoperonospora cubensis

16 Kingdom: FUNGI One cell or hyphae organisms with cell-wall containing chitin, flagella (only in chytrids) without mastigonemata, nutrition exclusively absorptive Phylum : CHYTRIDIOMYCOTA Unicellular, or primitive chain of cells, sexual reproduction by fusion of motile gametes; asexual reproduction: sporangium with uniflagellate zoospores Synchytrium endobioticum

17 Phylum : EUMYCOTA Hyphae or yeasts; absence of motile spores Subphylum: Zygomycotina Aseptate hyphae, sexual reproduction by fusion of sex organs (gametangia) leading to thick-walled resting spores (zygospores); asexual reproduction: sporangium producing non-motile spores. Rhizopus nigricans

18 Subphylum: Ascomycotina Hyphae with cross-wall (septa) or yeast, sexual reproduction by fusion of antheridium or spermatium with trichogyne on ascogon leading to development of an ascus with ascospores, asexual reproduction: conidias Morchella esculenta Boletus edulis Subphylum: Basidiomycotina Hyphae (with dolipore septum) or yeasts; asexual spores rare in most groups; sexual reproduction by fusion of compatible hyphae, leading ultimately to production of basidiospores on basidia, sometimes on or in large fruiting body

19 Subphylum: Deuteromycotina Hyphae (with septa) or yeasts; sexual reproduction absent, rare or unknown; asexual spores (conidia) formed in various ways Septoria apii

20 Important differences between the Oomycota and true fungi Zoospores Lysine biosynthesis Oomycota Biflagellate; one smooth and one with mastigonemata Via diaminopimelic acid True fungi Uniflagellate; one smooth in the Chytridiomycota Via α-aminoadipic acid Mitochondria Cristae tubular Cristae plate-like Wall polysaccharides Cellulose present; chitin also in some species No cellulose; chitin usually present Wall proteins Hydroxyproline present Proline present

21 Features of the main groups of fungi Group: Perforate septae +/- Asexual sporulation: Sexual sporulation: Lower fungi : Zygomycotina - non-motile sporangiospores zygospore Chytridiomycotina - motile zoospores oospore Higher fungi : Ascomycotina + conidiospores ascospore Basidiomycotina + rare basidiospore Deuteromycotina + conidiospores none

22 Other systems I. Gaumann, Archimycetes organisms without cell wall in trophic stage Phycomycetes place of karyogamy and meiosis is zygota - with zoospores (chytridiomycetes, oomycetes, plasmodiophoromycetes, hyphochytridiomycetes) - without zoospores (zygomycetes, trichomycetes) Ascomycetes place of karyogamy and meiosis is ascus Basidiomycetes place of karyogamy and meiosis is basidie heterobasidiomycetes, homobasidiomycetes Deuteromycetes only asexual stage

23 Other systems II. Alexopulos, 1966 LOWER FUNGI Myxomycetes Phycomycetes HIGHER FUNGI Ascomycetes Deuteromycetes Basidiomycetes Chytridiomycetes Oomycetes Zygomycetes Hemiascomycetes Euascomycetes Homobasidiomycetes Heterobasidiomycetes

24 Other systems III. Arx, Kreisel, 1969; Webster 1970; Ainsworth 1973 Myxomycota (Acrasiomycetes, Myxomycetes, Plasmodiophoromycetes, Labyrinthulomycetes, Trichomycetes) Chytridiomycota Oomycota Eumycota Zygomycetes Endomycetes Ascomycetes (Protoascomycetidae, Ascohymenomycetidae, Ascoloculomycetidae) Deuteromycetes Basidiomycetes (Holobasidiomycetidae Phragmobasidiomycetidae)

25 Other systems IV. Hawksworth, Kirk, Pergler, Sutton & Ainsworth, 1994 Myxomycota Dictyosteliomycetes Myxomycetes Plasmodiophoromycetes Eumycota Mastigomycotina Oomycetes Chytridiomycetes Zygomycotina Ascomycotina Basidiomycotina Deuteromycotina Basidiomycetes Ustomycetes Teliomycetes Coelomycetes Hyphomycetes

26 Other systems V. Cavalier-Smith, DOMAIN: EUKARYA 1. KINGDOM: PROTOZOA Acrasiomycota Myxomycota Plasmodiophoromycota 2. KINGDOM: CHROMISTA (STRAMENOPILA) Labyrinthulomycota Oomycota (Peronosporomycota) Hyphochytriomycota 3. KINGDOM: FUNGI Chytridiomycota Microsporidiomycota Zygomycota Ascomycota Basidiomycota

27 FOSSIL RECORDS ABOUR FUNGI The origin of the fungi appears to be very ancient They first appear in the fossil record coincidentally with the appearance of the land plants. Cambrium, Ordovician ( mil) - Marine Oomycota and Chytridiomycota associated with algae and free living Silurian ( mil) - Septate fungi in association with plants and cyanobacteria Devonian ( mil) - Lignolytic fungi, Zygomycotina and Basidiomycotina Carboniferous ( mil) - Basidiomycotina on woody plant residues Permian, Triassic ( mil) - Ectomycorrhizas formed with Basidiomycotina, Polyporales well established Jurassic, Cretaceous ( mil) - Ascomycotina increasingly evident Tertiary (10-75 mil) - Development of ericoid endomycorrhiza with Ascomycotina and possibly some Deuteromycotina and Basidiomycotina

28 Period Age (x 10 6 ) Major features of plant life in relation to fungi Possible features of fungi Tertiary Modern flora established Development of ericoid endomycorrhiza with Ascomycotina and possibly some Deuteromycotina and Basidiomycotina Cretaceous Conifers dominant, appearance of Pinus spp. and potentially mycorrhizal Angiosperms Ascomycotina increasingly evident Jurassic Luxuriant conifer and fern forests Ectomycorrhizas formed with Triassic Conifers Basidiomycotina, Polyporales well established Permian Conifers Carbonifer ous Early Gymnosperms Basidiomycotina on woody plant residues Devonian Vascular plants, rhynia Lignolytic fungi, Zygomycotina and Basidiomycotina Silurian Evidence for lichenized algae, a bryophyte-like land flora and emergence of vascular plants Ordovician 425 Septate fungi in association with plants and cyanobacteria Cambrian 500 Marine Oomycota and Chytridiomycota associated with algae and free living

29 MODEL OF RELATIONSHIPS BETWEEN THE THREE MODES OF FUNGAL NUTRITION Biotrophs fungi must obtain nutrition from living material (plant, animal) Necrotrophs - fungi occur on living material, but must kill part of tissue for obtaining nutrition (toxins) Saprotrophs - fungi living on dead material

30 Econutritional groups of fungi according to nutritional mode and ecological behaviour Obligate biotrophs (OB) Hemibiotrophs (HB) Facultatively saprotrophic hemibiotrophs (FSH) No capacity for saprotrophy or necrotrophy Initially biotrophic but then becoming necrotrophic, saprotrophic potential as for obligate necrotrophs Initially biotrophic but then becoming necrotrophic, a final saprotrophic phase then occurs Obligate necrotrophs (ON) Normally necrotrophic, any saprophitic ability severely limited or restricted to survival in dead tissues Facultatively saprotrophic necrotrophs (FSN) Facultatively necrotrophic saprotrophs (FNS) Obligate saprotrophs (OS) Facultatively biotrophic saprotrophs (FBS) Facultatively saprotrophic biotrophs (FSB) Normally necrotrophic with ability to become saprotrophic Normally saprotrophic with ability to become necrotrophic No capacity for necrotrophy or biotrophy Normally saprotrophic with some ability to become biotrophic Normally biotrophic with some ability to become saprotrophic

31 Concepts in fungal nutrition Nutritional behaviour Saprotrophic Necrotrophic Biotrophic Ecological behaviour Obligate Saprophytic Facultative Symbiotic Obligate Symbiotic = combination is creating parasitic form - = non-parasitic form

32 1. OBLIGATE BIOTROPHY Common features: Intracellular penetration by cells or haustoria Invagination of plasmalemma and following perforation Ektoparasites (Erysiphales), endoparasites (Peronosporales) Hypersensitivity Host nucleus is living Host specialization Complicated or impossible cultivation in axenic culture Examples: Plasmodiophora brassicae (Plasmodiophoromycota), Synchytrium endobioticum (Chytridiomycota), Peronosporales (Oomycota), Erysiphales, Taphrinales (Ascomycotina), Pucciniales, Ustilaginales, Tilletiales (Basidiomycotina) Blumeria graminis

33 2. HEMIBIOTROPHY Form between necrotrophy and biotrophy Cultivation in axenic culture possible Highly reduced saprotrophic existence Examples: Venturia inaequalis, Rhynchosporium secalis, Phytophthora infestans Phytophthora infestans Venturia inaequalis

34 3. FACULTATIVELY SAPROTROPHIC HEMIBIOTROPHY Form between biotrophy, necrotrophy and saprotrophy Cultivation in axenic culture possible Highly specific parasites Frequently occurring in 2 different forms : monokaryotic mycelium (biotrophic) and dikaryotic mycelium (saprotrophic) Crinipellis perniciosa (Basidiomycotina, obligate parazite of cacao)

35 4. OBLIGATE NECROTROPHY Not good developed ability for saprotrophic existence, time limited Persistence in death host tissues (mycelium, sclerotia, rhizomophs) Broad spectrum of hosts, diseases of fruits, leaves, stems and roots No problems with cultivation in axenic culture Sclerotinia sp. Hlízenka Armillaria mellea, Václavka obecná

36 5. FACULTATIVELY SAPROTROPHIC NECROTROPHS 6. FACULTATIVELY NECROTROPHIC SAPROTROPHS Ability of saproprophic and necrotrophic nutrition In contact with living host plant they are changing for necrotrophy After depletion of nutrition, persistence on death organic substrates, Broad spectrum of hosts, diseases of fruits and tubers No problems with cultivation in axenic culture Examples : Aspergillus, Penicillium (Ascomycotina) Rhizopus (Zygomycotina) Rhizoctonia, Botrytis (Deuteromycotina) Penicillium, Aspergillus

37 7. OBLIGATE SAPROTROPHY No ability of necrotophy or biotrophy Possibility of obtaining of nutrients from decomposition of sugars as well as more complicated compounds (cellulose, chitin, keratin) Onygena equina Panaeolus on cattle dung

38 8. FACULTATIVELY SAPROTROPHIC BIOTROPHY 9. FACULTATIVELY BIOTROPHIC SAPROTROPHS Ability of biotrophic and necrotrophic nutrition Frequently linked with occurrence of two different forms of pathogen: monokaryotic mycelium (saprotrofic phase) and dikaryotic mycelium (biotrofic phase) Highly specific parasites Examples: Hemiascomycetes Taphrinales (Ascomycotina), Taphrina deformans