MIcology

www.ipps2.blogfa.
com
2B
. .

. .
. .
Mycetology (mykes = mushrooms ) myceto
) logos (= discourse
.
Mycology .
.
) (
.

.
.
mushrooms
toadstools . ) (Cladius Caesar 54

Amanita phalloides .
15
) (Herbals ) ( .
) (G. E. Clausius 1601
Rariorium Plantarum Historia .
1675 ) (van Sterbeeck
) (Theatrum Fungorum
. . Leeuwen
) (hook .
) (spontaneous generation
www.ipps2.blogfa.com
. Micrographia )
( . Phragmidium Mucor
. .
(Piere Antonio
) Michelli
1729 Nova Plantarum Genera
.
Aspergillus, Mucor, Botrytis

.
Torurnefort ) Element de Botanique (1694 6 .
.

)(Carl V. Linnaeus
)(binomial nomenclature
Species
) Plantarum (1753 ) (Verms Chaos .

.
Baillard, Persoon, Link, Batasch, Schweinitz Fries

.
Synopsis Methodica Fungorum ) (Persoon 1801
) Systema Mycologicum (1821-1832 ) (Fries . 26

71 .
Leiden .
Gasteromycetes, Uredinales Ustilaginales ) (1821
. Schweinitz
. 4000
1200 . ) (Corda
6 Icones Fungorum Hucusque Cognitorum
. ) (Luis and Charles Tulasne
3 ) Selecta Fungorum Carpologia (1861-1865

) (polymorphism
.
) (Cooke
1872 Grevillea . ) (Buller

. ) (Research on Fungi 6
.
19 .
) (de Bary, 1831-1888
.
...
) (Puccinia graminis tritici 1886

1887
Comparative Morphology and Biology of Fungi,
Mycetozoa and Bacteria .
Phytophthora infestans
. ) (Woronin
) (Plasmodiophora brassicae
. ) (Brefeld
. ) (Saccardo
25
Sylloge Fungorum Omnium Hacusque
Cognitorum 1882-1931 .
1880 .
19 .

.
.
. .
) van Tieghem (Raulin
) () Wehmer (
. .
Aspergillus niger
. .
20
.
80000
-120000
1/5
.
.
.
.
0B

chemoorganotroph

. )
(
.
] .
[.
) (soma
) (hypha .
)] .(totipotency soma
somatic vegetative
vegetative [ . thallus
.
.
.

.
.

.
.
.
) (Penicillium ) (Aspergillus
.
.
.
) (substrates
.
pH.

.
.

.
osmophilic fungi .
.
.
25-30
10 40 . ) (mesophilic fungi .
) (thermophilic fungi 40
20 . 50
thermotolerant ) Mucor pusillus,
Cheatomium thermophilum, Thermopascus auranticus .(Aspergillus fumigatus
) (psychrophilic fungi 20 ]
) Typhula spp.

Thamnidium elegans, Cladosporium
herbarum
Pythium
[ .
psychrotolerant .
.
62-65
4-7 .

pH . acid tolerant
)
Aspergillus, Penicillium, Fusarium 2 5/5-6
( .
) (acidophilic
1/25 .
Acontium velatum
pH 10-11
.Fusarium oxysporum, Penicillium variable pH pH

/2-0/3 0
) (
) (polyols .
.
Saccharomyces cerevisiae
. 4 :
-
) (obligate aerobes
.Rhodotorula, Phycomyces
-
) (facultative aerobes
) (
.Mucor hiemalis, Aspergillus fumigatus, F. oxysporum
-
) (obligate fermentative
Aqualinderella fermentans
Oomycota
Blastocladiella ramosa
Chytridiomycota . )(
)( .
-
) (obligate anaerobes
) (
) Neocallimasticales .(Neocallimastix )
(
(mixed acid
) fermentation .
]
.Cylindrocarpon tonkiense, F. oxysporum Aqualinderella CO2
5-20 . Coccidioides immitis CO2
[.

.
. .
.
) (saprobes .
) (parasite
.
.
) (obligate parasites=biotrophs
.
) (facultative parasites .
) (obligate saprophytes
.Trichoderma viride
) (facultative saprophytes .

.
) (symbiosis
) (lichens - )( ) (mycorrhizae
.

.
)(endophyte

.
.

.
)( .
. Lolium
perenne, Festuca arundinacea
.
) ( .
) ( .

. ) Laboulbeniales
Zygomycota (] .
[ .

) .(Trichomycetes
.
)(
.
) (mycelium .
) (yeast
.
) (dimorphic fungi .
. .
.
.
) (septum .
) ( .

) (monokaryotic
) (dikaryotic
) (polykaryotic . septate hyphae .

. a
) septate (= non septate coenocytic hyphae .

.
.
.

) ( .

) (woronin bodies .
.
) (dolipore septum .

) (perforate ) (imperforate .
) septal pore cap (SPC
parenthosome .

micropore
. Saccharomycetales
Geotrichum
] Candida albicans 25
[ .
homokaryotic
heterokaryotic . )
( heteroplasmon ] .
: ) (primary septum = eu septum

= (adventitious septum
) distoseptum
[.
.
.
.

.
) (
. .

. .

.
1-4
.
.
1-3 1-6 .
. )(
.
1-3 1-4
.
) ( .
.
. )
( .
.
10
1 4 .

1 4 .
) (polyuronoids
) (.
]
.
13 16.
, 14 [.
) (
. osmoenzymes

. .
.
. ATPase Mg-ATP
ADP . 3 1 1
4 .
.
.
.
.

azoles .allylamine
Pythium . Phytophthora
.
) (tip growth
. .
11
) (extension zone ) (Neurospora crassa

40 .
.
.
)(
) 100( )
100( .

spitzenkrper = apical body .

.
) (secretory vesicles
.

.
) (chitosomes .
Spitzenkrper
.
.
) (

) (fungal fimbriae
30 7 .
.
) (
) (extracellular matrix . .

.
.

.
. Hysterangium crassum
12
Sclerotium rolfsii
.

.

.
.

. plectenchyma . :
prosenchyma .
pseudoparenchyma
.
.
)(modification of hyphal structures
( )(rhizomorphs

) (
.
.
. .
) 3-4 (
vessel hyphae .
.
.
.
) (Armillaria mellea . 5-6 .
13
1 1000
. .
Marasmius
.
( )(mycelial strands

.
.
.
.

Phalllus impedicus
) (Serpula lacrymans .
.

) Agaricus bisporus (.
( )(appressorium
) (

. ) ( .
) = (penetration peg
.
) (Magnaporthe grisea.
( )(haustorium
) (
.

.
14
.
.
( )(traps

. .
-1 : -2

Stylopage grandis
Meria
Nematoctonus .Harposporium
Arthrobotrys oligospora .

.
Dactylella Zoophagus insidians .

. Dactylaria candidas D. gracilis .
( )(sclerotia

.
.
. Polyporus mylittae
15 . :
cortex, rind medulla
Sclerotium, Sclerotinia . ) (pseudosclerotium
Rhizoctonia .
.
Claviceps Sclerotinia sclerotiorum
] .[carpogenic germination
15
.[myceliogenic germination ] Rhizoctonia Botrytis cinerea

.
( )(hyphopodia
:
-
capitate hyphopodium
Gaeumannomyces .
-
mucronate hyphopodium Meliolales
.
}
. .
-1 ) (pertotroph or necrotroph

-2 . ) (biotrophs
. )(
-3 . ) (hemibiotrophs
) (
) Colletotrichum lindemuthianum Cercosporidium personatum
(.
)(Fungal Organells
)(nucleus

.

.

.
.
16
] ) [spindle pole bodies (SPB SPB .

.
. SPB .
SPB
)( . SPB
. SPB
.
. SPB
) (astral SPB
. SPB . )(
SPB .
) SPB
Centrosome Spindle plaque, Nucleus associated organelle
Microtubule organizing centre
. . SPB
) (
(.
) (intranuclear

. Dictyostelium discoideum
.

. ) (DAPI .
.
:

. .

pulse field gel
) electrophoresis (PFGE .
17
.
UV .
1B
)(mitochondria

.
.

. 1-20 DNA
) Hansenula mralcie ( .
ATP .
) (ribosomes
.
80S) . 70 S(.
) (vacoules
.
ATPase
.
) .

(.
) (golgi bodies

.
. .
) (lomasomes

18

.
) (hydrogenosomes
0/5-2 .
DNA .
) (chitosomes
.
40 - 70
1/3 . 2/3
.
) (rumposome ) (gama Particles .
Blastocladiella emersonii 0/5
. .

Monoblepharella
.
.
) (filasomes

.
.
) Multivesicular bodies (MVBs

MVBs SPB .
) (
.
. .
. SPB
.
.
19
)(Fungal Nutrition

.
: )(microelement= trace element

.
) (macroelements . :
.

. ) (boron
.
.
0/001-0/05 .
(yeast
) extracts .
50
.
.
.
.
. .

Leptomitus lacteus, Sclerotium rolfsii, Fusarium
.lini
catabolite
repression
. .
Verticillium .

.
. .
Sordaria fimicola
Rhizopus oligospora .
20
.
.
Leptomitus lacteus
.
. )
( ) (
. .
. .

.
) (casein hydrolysate
.
.
euthiotrophic .
parathiotrophic
.
. .
. .
. KH2PO4 K2HPO4
.
.
. 3+
2+ . .

. :
auxoautotrophic

Ashbya gossypii Eremothecium ashbyii .
auxoheterotrophic .

21
Pythium butleri . .
)
) (A, D, E, K .
B (H
Phytophthora, Phycomyces
Polyporus, Marasmius, Exobasidium .

.
Sordaria fimicola
. Glomerella cingulata
.
.
Polyporus texanus . .
.
5 : ) (lag phase
. ) (logarithmic= exponential
. ) (decceleration
. ) (stationary phase )(autolysis
.
. .
)(Reproduction and Sporulation
.
.
.
-1 : ) (plasmogamy -2 ) (karyogamy
-3 ) (meiosis .

)(meiospores
.
.
. )
(
. ) (holocarpic .
22

. ) (eucarpic .
.
) ( .
) (gametangium .

.
) (isogametangium

) (heterogametangium .
) (antheridium
)] (ascogonium ) [(oogonium .
.

.

.
) (mitospores .
:
-1 ) (fragmentation .

.
) (arthrospores
Geotrichum .
.

. ) (chlamydospore.
-2 ) (fission
) (
) (Schizosaccharomyces.
23
-3 ) (budding .
.

. ) (blastospores .

) (pseudomycellium . .
-4
.
.
) (sporangium
)(sporangiospores
. ) (sporangiophore
.
aplanospore.
) (zoospore or planospore
. .
) (whiplash . ) (tinsel
. .
)
) (heterokontean type( .
) 2
( . ) (conidium .

) (conidiophores . diaspore

) ... ) (diaspore(.
) (parasexual cycle .

Pontecorvo and Roper
) (1952 .
24
.
:
- 1
) (heterokaryosis

.
- 2
- 3
) (
- 4
) (sectors
.

2N+1 2N-1 . ) (aneuploids . 2N-1
.
.

.

. .
) (cytoplasmic inheritance .

.
.

.

.
. barrage phenomenon
.
25

) (zone line .
.
)(sex and sexual compatibility

.

.
) (mating types .

) ( .
.
:
-1 ) (hermaphrodite or monoecious
.
-2 ) (dioecious .
. .
-3 ) (sexually undifferentiated
.
.
:
( ) :(homothallic fungi
.
mating type )
(.
( ) (heterothallic fungi
.
.
- ):(secondarily homothallic fungi
26

) (
.
) (pseudohomothallism Neurospora tetrasperma Agaricus

bisporus .Exobasidium japonicum
:
-1 ) (
.
-2
)
(:
(I
) :(bipolar or unifactorial
) ( a + A
Mucor, Rhizopus, Phycomyces ) a A
(.
(II
) :(bifactorial or tetrapolar

Schizophyllum commune . A B
2 .
Schizophyllum commune A B
.
A .
A
.
flat reaction .
) (
. . barrage reaction.
) (teleomorph
) (anamorph . mitosporic fungi meiosporic fungi
27
. ) (holomorph

andromorph . .

synanamorphs.
1881 Achlya .
erotactin .
erogen
erotropin . :
Trisporic acid Antheridiol Sirenin .
Sirenin Allomyces ) (
Trisporic acid
) Antheridiol .(erotactin
erogen .
erotropin Achlya .
Blakeslea trispora Mucor mucedo Phycomyces blakesleeanus
. .
a ) (

DNA ) sporopolenin .(
.
)(fungal life cycle
) (n ) (n+n ) (2n

.
-1
.
-2
Blastocladiales .
28
-3

. .
-4 -

.
-5 -

Endomycetales ) Blastocladiales
Allomyces,
(Coelomomyces ) alternation of generation (.

-6
.
] diplomitotic nuclear cycle

: .
haplomitotic A nuclear cycle :

.
haplomitotic B nuclear cycle

[.
)(Importance of the Fungi
250
. .
Osiris .

Jupiter
.
Amanita muscaria .
) (hallucinogenic Psilocybe cubensis
. ) = (bioluminescense .

.
29
.
.

.
) (fox fire . Armillaria bulbosa
) (1992 .
30 10
1500 .

.

) (
.
) ( .

.
.
Serpula lacrimans
.

) (
.

.

) (mycotoxin
.

. .
30
ochratoxins Aspergillus ochraceous ) Penicillium viridicatum

(
. A, B C
B aflatoxins . Aspergillus flavus A.
) parasiticus ( ) Penicillium
(

.
fumonisins Fusarium moniliforme
zearalenones .
) 12-14
( vapona .
patulin Aspergillus Penicillium )

( . .
trichothecenes
Fusarium, Trichoderma,
Stachybotrys, Myrothecium, Cephalosporium Trichothecium roseum

. ) (penicillic acid
Penicillium puberulum Aspergillus ochraceous ) (
. ) (sporodesmin Pithomyces chartarum

.
8 . 3 G, B
M . B UV G
UV . M
. B1 DNA
.
Flavobacterium .
31
Claviceps purpurea
.
) (ergot .

.

) (St. Anthonys fire or holly fire
.

14-18
.
) (ergotism . .
-1
) (lysergic acid dimethylamide = LSD -2 ) .(clavines

.

.
. -
. Amanita
:
phallotoxins
amatoxins 10-20 .
phalloidin
.
.
Lepiota Galerina
orellanine . Cortinarius orellanus
) 3-15( .
Amanita
muscaria Clitocybe napipes

.
psilocybin psilocin
Psilocybe,
Stropharia, Coenocybe
.
32
.

.

] ) (Ophiostoma novo-ulmi
10 [
]) (Cryphonectria parasitica [.
) (mycosis .
.
.

.
.
) (blastomycosis ) (coccidiosis ) (histoplasmosis
) (candidiasis ) (crypyococcis ... Candida albicans .

.

7-8 .
.
3B

.
.
Penicillium chrysogenum 1928 .
Cephalosporium acremonium

. sandimmune sandimman

.
1970
33
Cylindrocarpon lucidum
Tolypocladium inflatum .
Taxol .
Fommitopsis officinalis
.
Claviceps purpurea

. )(
. .

... .
600
. 1650 .

. .

Agaricus brunnescense .
. ) (
Agaricus brunnescense .
)(aphrodisiacs
.

) (truffles
) (morels .

. ) (Ustilago maydis

huitlacoche cuitlacoche .
Brie, Commembert, Roquefort Danish
Blue . .
miso tempeh sufu
.

34

.
CO2 .

.
(malting) .

. B .
.
. ) (mycoherbicides Colletotrichum
gloeosporioides Collego
) Aeschynomene virginica ( .
.
.
.
.

.
Ambrosia

. mycangium
.
.

anobiid .
.

.

35
. Neurospora, Saccharomyces,
Schizophyllum ....
)(Fungal Systematics
3 5

.

: ) (
) (

) (kingdom Plantae ) (kingdom Animalia
.

.
1860
) (Hogg Protoctista . 1956 ) (Copland
) (Monera 1969 ) (Whittaker
) ( .
: ) (Kingdom Monera
) (
) (Kingdom Protista

. ) (Kingdom Plantae
. ) (Kingdom Animalia
) (Kingdom Fungi
.

.

36
.
) (1993
Cavalier-Smith
Eubacteria, Archaebacteria, Archezoa, Protozoa,
Plantae, Chromista, Animalia Fungi

) (rDNA .
) taxonomy ( taxo
nomy .

.
. :
) (classical taxonomy .
) (biochemical .
) (numerical . ) (molecular
. ) (cytotaxonomy
.
) nomenclature(
) identification .(
) systematics .(
) ontogeny .(
.
) phyllogeny(

.

.
) monophyletic (
) polyphyletic . (
. .
paraphyletic ) ( .
Fungi
. .
37
phylogenetic
classification ) (
.

. ) (character
. ) (character states.
)
( .
.
. 1887
)( . )
( .
.
.
1960 .

.
.
.
.

. .
.
.
.
RNA
.rDNA
38
rRNA
( . rDNA
.
RNA) (18S .
.
.
. : plesiomorphs
.apomorphs
symplesiomorphs
synapomorphs . .
.
.
) (sister groups
. .
:
.
. .
maximum parsimony, maximum likelihood distance .

. distance phenetic
.
.
)
(
) /
( .
DNA
. .
maximum parsimony ) (
.
.
39
.
.
. .
.

. maximum likelihood
.
. maximum likelihood

.
.
.

maximum parsimony .

) (robustness .

.
.
) (taxon ) ( . .
) (binomial
) (species epithet .
.

. .

.
.

40
:
. :
.
: ) (-mycota ) (-mycotina ) (-mycetes ) (-mycetidae (-
) ales ) (-ineae ) (-aceae ) (-oideae ) (-eae ).(-inae
.

) 5 (
Chromista, Fungi Protozoa .

2007 Fungi
Chytridiomycota, Zygomycota, Glomeromycota, Ascomycota Basidiomycota

. Chromista Stramenopila
Hyphochytriomycota Oomycota, Labyrinthulomycota .
Myxomycota, Acrasiomycota, Plasmodiophoromycota Dictyosteliomycota
) Protozoa (Protista
. 2007

.
7 Neocallimastigomycota
Basidiomycota, Ascomycota, Glomeromycota, Chytridiomycota, Blastocladiomycota,
Microsporidia Zygomycota

. :
Mucoromycotina, Entomophthoromycotina,
Zoopagomycotina .Kickxellomycotina
incertae sedes .
.
.
.
41
)(International Code of Botanical Nomenclature = ICBN

4
. 6 62
5 .
:
-1 ) (priority .

.
.

.
)
(spec. nov.
) 1935
(.
-2 ) (typification )
(
. .
. )
DNA . 1993
(.
) (nomenclatural types :
) (holotype
.
) (isotype
.
) (lectotype

.
42
)(paratype

) ( .
)(neotype
.

) (epitype )
1958 (.
-3 ) (authority
.
. Mucor corymbifer Cohn 1884 Cohn
. 1912
Absidia
. :
Absidia corymbifer (Cohn) Saccardo & Trotter

) ( )
(basionym
)
.(1953 ) (1958

.
Schiffnerulla
pulchra (Sacc.) Petrak Sarcinella heterospora Sacc.

Guestieriella pulchra S. Hughes .

DNA
. Taxon
.
) (homonym .
) (synonym
.
) (legitimate name .
) (correct name
.
43
)(Ascomycota

) (ascus .
) (
. )(free cell formation
. .
) (woronin bodies .
.

)
( .

) (coprophilous fungi .
) (corticolous
) (lignicolous ) (folicolous .
hypogeous )
epigeous( . -
)( )( .
) (mycobiont ) (photobiont 46 .
96 .
.

. .
)
Trichoderma (Chaetomium
. ) (
.
.
Beauveria, Fusarium,
Metarrhizium
. Ambrosiella, Symbiotaphrina, Raffaelea
.
.
.
44
Gibberella
) fujikuroi ( . )
(
. Neurospora crassa, Saccharomyces
Aspergillus .

. .
.
(fungi
) imperfecti .
.
:
.
.
DBB (diazonium blue
) B .
) hologenous
( ) entrogenous
( . - ) (%G+C DNA
50 ) (50-70 50 ) (28-50 .
.

.

. .

.
.
45
)
( .

.
.
.

) (soredium
. .
):(sexual reproduction
:
- 1 ) (gametangial contact .
) ( .
) (trichogene
)( . .
.

.

)(conjugate divisions
. .
)
crozier
( .
.

. .
.
(ascus mother
) cell .
.
.
4 .
46
8 .
ascus vesicle envelope membrane
) system (EMS .
.
. ) (sporoplasm
) (epiplasm .
.
8 1000
. ) (
EMS} .
{.
- 2 ) (gametangial copulation hologamy

.
.
- 3 ) (spermatization deutrogamy

.
.
) (spermogonium .
.
- 4 ) (somatogamy
. .
) (.
47
- 5 ) (apomixis
.
.Ascobolus aquinus
)(asci
) (
. )(Hansenula anomala

) (Endomycopsis fibuliger
) (Emericella variecolor .
) (hymenium .
. ) (protounicate
.
) (passive . ) (eutunicate
. :
) (unitunicate
) (operculum
.
) (bitunicate
) (exotunicae ) (endotunicae
.
. fissitunicate jack in the
box . ) (active.
)(ascocarp = ascoma

.
. :
) (cleistothecium . ) (perithecium
-. ) (apothecium
.
. ) (ascostroma
48
.
.

) (pseudoperithecium -. ) (thyrothecium
. ) (hysterothecium
.
.

) (hamathecium . :
- 1
) (interascal pseudoparenchyma
- 2
) (paraphyses
.Wettsteinia

.
.Xylaria, Pyrenula
) (epithecium
Genea .Pezizales
- 3
) (periphyses
Gibberella,
.Pyrenula
- 4
) (apical paraphyses
) (Periphysoids
.Nectria, Metacapnodium
- 5
) (pseudoparaphyses

.
.Pleospora
49
- 6
trabeculate pseudoparaphyses
.
- 7
.Dothidea .
)(incompatibility systems in ascomycota

. .

.
.
homogenic incompatibility .

) (mating type genes . heterogenic compatibility
.
(heterogenic
) incompatibility genes
(heterogenic
) incompatibility .
.
) (vegetative compatibility groups .
mating type .
) (idiomorph
.
)(ascomycota classification

.
Dictionary of Fungi 46264
3226 32267 Alexopoulos ) (1996

:
.
6 :
50
Pyrenomycetes -1 .
Discomycetes -2 .
Loculoascomycetes -3 .
Laboulbeniomycetes -4
) (cladome.
Plectomycetes -5
.
Hemiascomycetes -6 .
.
:
-- .
-1 - -2
Endomycetes
Ascomycetes .

2007
Taphrinomycotina, Saccharomycotina Pezizomycotina

) = (asexual fungi

.
.
.

.
.

.
.
) (conidiomata
. :
-
) (synnemata= coremium
51
.
.
-
) (sporodochium

.
-
) (pycnidium
.
.
.
.
.
-
) (acervulus
.
.

) (conidiogenous cell .
.

.
)
= (muriform .

. macrocyclic conidiation .

. microcyclic conidiation .

( .
)(conidial ontogeny
52
) Zygosporium
:
- a
) (blastic

.
- b
) (thallic
.
- c
) (thalloblastic
.
- d
) (arthric
.
.

hologenous .
entrogenous .
endogenous .
) (solitary .
) (catenate seriate .
. seriate
) (false chains
. (spore
) balls
.
basipetal
acropetal
. :
-
schizolytic
.
-
rhexolytic
.
.
53
Onygenales
36B

. .
) (form .
.
Sphaeropsidales Melanconiales . .
Moniliales
. ) (mycelia sterilia
) ( .

) (sections
. :
Amerosporae -1 .
-
Allantosporae ) ( .
Hyalosporae .
Phaeosporae .
Didymosporae -2 .
-
Hyalodidymae .
Phaeodidymae .
Phragmosporae -3 2 .
-
Hyalophragmiae .
Phaeophragmiae .
Dictyosporae -4 .
-
Hyalodictyae .
Phaeodictyae .
Scolecosporae -5 .
Staurosporae -6 .
54
Helicosporae -7 .

) :(Barron, 1968
-
Arthrosporae
.Geotrichum
-
Blastosporae .
.Cladosporium
-
(Radulosporae) Sympodulosporae

.Arthrobotrys, Sporothrix
-
Porosporae
.Alternaria, Bipolaris, Curvularia

-
Aleuriosporae
. .
.Histoplasma, Blastomyces
-
Annelosporae
Spilocea, Scopulariopsis
-
Phialosporae
. basipetal .Aspergillus, Penicillium, Verticillium

-
Botryoblastosporae
. )
(Botrytis
) (Oedocephalum .
-
Meristem Arthrosporae basipetal
) Acrosporium (Oidium
.Trichothecium roseum
-
Meristem Blastosporae
.Arthrinium phaeospermum
.
55
:Hyphomycetes -1 . .
) ( .
.
:Coelomycetes -2 .
.
:Blastomycetes -3 .
.
.
Aganomycetales .
:
Rhizoctonia .
.

.
. .
.
.
Sclerotium .
.
Sclerotium rolfsii .
. ) Botryobasidium rolfsii (
.
.
.
Sclerotium cepivorum
Sclerotium oryzae )(
) Magnaporthe salvini ( Nakataeae sigmoidea.

(Hyphomycetales) Moniliales
56
micronematous ) (
) macronematous ( .
.
. :
Moniliaceae
) ( .
.
. :
Aspergillus
.
.
. )(
.
.
Penicillium
.
) (monoverticillate metula
) .(biverticillate .
. P. notatum P. chrysogenum
P. griseofulvum
. P.
expansum P. digitatum P. italicum
)( P. gladioli . .
Pyricularia
1-3
. P. grisea Magnaporthe grisea.
Geotrichum
57

.
.

. .
Geotrichum candidum
Endomyces geotrichum . .
Verticillium
.
) .(verticillate .
. V. dahliae V. albo-
atrum .
V. lecanii

Lecanicillium lecanii .
Trichoderma
.
.
.
. Hypocrea .
T. virens viridin gliotoxin
4B
.
Clonastachys
Gliocladium .
penicillate .
.
.
58
C.
rosea .
Dematiaceae
.
.
Alternaria
.
.
.
. .
.
) beak(
. Lewia .
Curvularia
.
3-4
porosporae.
. C. lunata Cochliobolus lunata

.
5B
Cercospora
.
.
. .
C. beticola . C. arachidicola
C. musae )( .
Mycosphaerella
musicola C. apii .
.
59
6B
Botrytis

)( .
. B. cinerea
B. allii
B. fabae .
Botryotinia .
Drechslera

porosporae.
.
Cochliobolus Pyrenophora .
Stilbellales
. .
Stilbellaceae . Graphium Trichurus.
Graphium
.
annellidic .
Ophiostoma .
Tuberculariales

. Tuberculariaceae .
.
Fusarium, Tubercularia, Epicoccum
Exosporium.
Fusarium
. . :
.
.
60
.
. Nectria, Gibberella, Calonectria
Hypocreales . .
Tubercularia
.
T. vulgaris . Nectria cinnabarina
.
Coelomycetes
. 9000

.
.
Sphaeropsidales,
Melanconiales Pycnothyriales .
Sphaeropsidales
.
. . 4
.
Sphaeropsidaceae
.
Nectrioidaceae .
Leptostromataceae .
17B
16B
Excipulaceae .
Sphaeropsidaceae :
Phoma
.
.
. .
61
Pleospora
. P. glomerata
.
) P. betae (tel. Pleospora bjorlingii
P. lingam (tel. Leptosphaeria maculans) .
Phyllosticta
.
.
. .
Guignardia Mycosphaerella . P. Citricarpa .
Macrophomina
.
. 3-4 .
.
. M. phaseolina
. .
Phomopsis
.
stylospore
. .
Diaporthe . P. cinerescense P. vexans
P. mali P. leptostrmiformis .
Ascochyta
.
. .
. A. rabiei A. fabae
A. phaseolorum .
Leptosphaeria .
Didymella
62
Mycosphaerella Didymella
Ascochyta .
Septoria
.
. .
S.
apiicola S. tritici S. pyricola (tel. Mycosphaerella

) sentina . Leptosphaeria Mycosphaerella
.
Nectrioidaceae
.
Aschersonia
.
.
. .
Hypocrella .
Melanconiales
Melanconiaceae .
. . .
7B
Colletotrichum

. .
setae .
.
C. gloeosporioides
Glomerella cingulata
.
Glomerella .
C. lindemuthianum
C.
) orbiculare (syn. C. lagenarium C. coccedes

.
Marssonina
63
.
. M. rosae M. juglandis
)( . Diplocarpon .
Pestalotiopsis
5 .
.
. P. theae .
Entomosporium
18B
.
.
E. maculatum . Diplocarpon mespili
.
Blastomycetes
.
:
Cryptococcales
Cryptococcaceae
. . .
.
Cryptococcus, Candida,
Trichosporon, Malassezia, Rhodotorula Torulopsis . Rhodotorula

.
R. glutinis
. .
Rhodosporidium Ustilaginsles.
19B
Candida .
. Candida Leucosporidium .
Cryptococcus .
C. albidus
floriforme C. neoformans F. neoformans.

Sporobolomycetales
64
Filobasidium
4 ) (mirror yeasts

.
. .
Sporobolomyces, Bullera, Itersonilia Tilletiopsis .
(Archiascomycetes) Taphrinomycotina

39B
rDNA .
)(
.

.
Schizosaccharomycetes, Neolectomycetes, Pneumocystidomycetes Taphrinomycetes

. Saitoella ) .(incertae sedis
Neolecta rDNA
(1 :
Saitoella (2 .
(3 . (4 .
.
Pneumocystis Pneumocystideaceae Pneumocystideales
3 )( .
.
1988 ) nSSU-
(rRNA .
) P. carini (
)-(
P. jirovecii
.
Neolecta Neolectaceae Neolectales 3 .

.
65
EMS ) (.
.
.
Saitoella S. complicata
Rhodotorula glutinis
. .
. )
( ) ( .

.
66
Taphrinales Taphrinaceae Protomycetaceae

.
Taphrinaceae
)( .

.
. Taphrinaceae Taphrina 95
)
( .
T. deformans
. )
( .
Taphrina Lalaria .
.
.
.

.
.
.
.
(ascus
) mother cell 8 EMS

8 . (foot
67
) cell
.
.
. .
T.
pruni T. cerasi T. coerulescense

T. bullata T. betulina T. populina
T. insititiae T. wiesneri .
T. epiphylla .
Exobasidiales Ustilaginales )
(.
Protomycetaceae 5 20 Protomyces

) Compositae (Asteraceae
) Umbelliferae (Apiaceae . Taphridium Burenia Umbelliferae

Protomycopsis Volkartia Compositae .
.
:P. inundatus
.
. .
.
68
.
.
.
. .
.
) (synascus .
.
.
Schizosaccharomycetales
. S. pombe
.
S. octosporus S. japonicus
. .
S. japonicus
.
.
.
.
.
.
.
8 8 .
. S. octosporus
S. pombe .
Saccharomycotina )(the ascomycetous yeasts

1000 .
69
.
.
.
( .
)( 10 .
) (
. ) (

.

.
.

)( . .
.
.
.
. .

) (ascophore
EMS .Cephaloascus
.
1 3 1 6
. rDNA
.
polyoles
.
.
)
(killer toxins
70
Saccharomyces
cerevisiae .

dsRNA
Pichia, Candida, Kluyveromyces, Ustilago maydis
Cryphonectria
parasitica .

.

.
DNA .

GC DNA .
28-50 50-70 . 1-2

.
DNA ) DNA
( . 80 .

70
.
DNA .
Saccharomycetes Saccharomycetales 13
).(incertae sedis

Ascoideaceae, Cephaloascaceae, Dipodascaceae,
Eremotheciaceae,
Lipomycetaceae,
Metschnikowiaceae,
Saccharomycodaceae,
Saccharomycopsidaceae,
Trichomonascaceae, Saccharomycetaceae.
Endomycetaceae,
Phaffomycetaceae,
Saccharomycetaceae
:
.
.
.
cerevisiae .
Saccharomyces
Kluyveromyces, Zygosaccharomyces, Torulaspora,
Kazachstania ... . S. cerevisiae
71
a .

.

.
sexual agglutination
. .

. .
4 4 .
.
. Kluyveromyces

) .
Saccharomyces Dekkera Kluyveromyces
( . K. polysporus
.
Saccharomycodaceae
) (apiculate
. . Saccharomycodes ludwigii
.
) A1 (A2
.
) (sprout mycellium
. .

4 .
.
Eremotheciaceae
72
Saccharomycetales
)(Hemiptera
) stigmatomycose
( . .
Eremothecium Coccidiascus .
1-2
.
Eremothecium (Spermophthora) gossypi
.
.
. . 40
) ( . )(

) (2N
. .
8

Cephaloascaceae

Cephaloascus Ambrosiozyma .
.
. C. fragrans
.

.
Sporothrix
Ophiostoma . .
.
. .

73
.
4 .
Lipomycetaceae

. Lipomyces
Zygozyma ) Dipodascopsis (
. Dipodascopsis uninucleatus
.
. )(
.
.
.
.
Dipodascaceae
.
DNA
.
Dipodascaceae
Dipodascus Galactomyces Geotrichum
.
. Dipodascus albidus
.
) (
.
.

.
74
.
.
Ascoidea Ascoideaceae
.
.
.
16-160 .
) (A. asiatica
) (A. rubescens .
.
Metschnikowia Metschnikowiaceae
1-2
.
.
.
. .
1-4
8
1-2 . . 8
)(Co-Q8 . .
) Pezizomycotina = (Filamentous Ascomycetes

.
.

.
.
75

. Nonfeldt 1932 3
Ascohymeniales
Plectoascales :
Ascoloculares . Luttrell 1951 ) (centrum

Plectoascomycetidae
Hymenoascomycetidae
Loculoascomycetidae
Laboulbeniomycetidae
.
.
(Eurotiales and related species) Plectomycetes
:
. ) ( )
(
reticuloperidium .
. )
8 ( .
.
. ) (
. ) ( Eurotiales

.
. rhexolytic Onygenales Ascosphaerales.
schizolytic
76
Plectomycetes .

.

.
37
.

. Ascosphaerales, Onygenales Eurotiales
.
Ascosphaerales
4 1 ) Arrhenosphaera( 11 ) Ascosphaera (
1 ) Bettisa( 2 ) Eremascus( 3 .
.
. xerophilic 0/85
60 .
DNA . Ascosphaera
. .
Ascosphaera apis
) (chalkbrood.

.
.
30-40 .
Ascosphaera
.

.
.
) (cyst
nutriocyte .
) ( .
77
.
A. atra . .
Erymascus
. . E. fertilis
.

rhexolytic .
)) (copulation(
.
. 8 8 .
Onygenales
:
-
.
.
. .
rhexolytic -.

.
Onygenaceae, Myxotrichaceae, Gymnoascaceae Arthrodermataceae
. .
Onygenaceae ) (
. .
.
.
Ajellomyces .
) Ajellomyces dermatitidis (Blastomyces dermatitidis ) A. capsulatus

(Histoplasma capsulatum
37 .
.
78
Chrysosporium .
. Onygena eqina
.
mazaedium
)
( . O. corvina
.
} geophilic .
zoophilic anthropophilic .
{.
Eurotiales
:
- ]
) (peridium
[ .
. .
. schizolytic
. .
Trichocomaceae Pseudourotiaceae .
Aspergillaceae) Trichocomaceae (Eurotiaceae

Aspergillus, Penicillium .Paecilomyces Monascus

Basipetospora .
.
. Penicillium roqueforti P. commemberti
.
Penicillium
notatum P. chrysogenum P. griseofulvum

.
79
Aspergillus fumigatus .
.
A. niger

. .
2/5 10-9
.
: Talaromyces Byssochlamys
.
. .
Penicillium Paecillomyces
) (biverticillate . Eurotium, Emericella, Eupenicillium, Neosartorya

Hemicarpenteles
.
. .

) ] .(bivalvate
Paecilomyces Byssochlamys Aspergillus 9

Eurotium, Sartorya Emericella . Penicillium
Talaromyces, Eupenicillium Carpenteles[.

.
.
. .

Aspergillus
. (foot
) cell .
. .
biseriate
metula .
. uniseriate
80
. basipetal .
.
.
. .
.
.
.
... .
Penicillium Aspergillus .
2/3
.
penicillus .
.
.
.
Penicillium
.

) Eurotium
Aspergillus glaucus (

.
.
.
.
)
( .
.

.
.
Monascus
81
.
.
. .
Basipetospora .
Emericella .
hlle cells
.
. .
diaspore .
.

) Aspergillus nidulans (.
Sartorya
. 8
Aspergillus
fumigatus .
) Paecillomyces Byssochlamys Aspergillus
11
Sartorya, Eurotium Emericella .
Penicillium Eupenicillium, Talaromyces Carpenteles.

Hemicarpenteles Eupenicillium

8 . Eupenicillium
.
.
.
. bivalvate
. Byssochlamys
.
Talaromyces flavus
Penicillium vermiculatus
.
82
. .

.
.

)(Pyrenomycetes, ascomycetes with perithecia
:
subiculum .

.
.
.
. schizogenous
. lysigenous Loculoascomycetes
) ( .
.
.

) (
.
Neurospora
. .
.

.
9 :
Hypocreales, Melanosporales,
Microascales, Phyllachorales, Ophiostomatales, Diaporthales, Xylariales, Sordariales,

.Meliolales
83
) Luttrell (1951 ) (
.
. Nectria
.
Hypomyces Hypocrea Nectria
.
Claviceps
) (
.
Hypocreales
-: - . )
( .
.

part spore
. .
.Hypocreaceae, Nectriaceae, Clavicipitaceae :
Hypocreaceae
Hypomyces, Hypocrea Sphaerostilbella
Hypocrea .
.
8 . .

16 .
Trichoderma ) Clonostachys (= Gliocladium .

Hypomyces subiculum
subiculum . )(
. 1-2 ) (
. Sepedonicum .
84
Nectriaceae

.
Giberella Nectria . Nectria
.
.
. 1-2 N. galligena .

. N. cinnabarina
.
.
.
.
Tubercularia -
.
.

.
. .

. 8
. .
N. haematococca
) ( .
Giberella
) G. zeae . (Fusarium graminearum
G.
xylarioides
) G. fujikuroii (Fusarium moniliformis )(
85
.
...
Clavicipitaceae

Hypocreales .
subiculum
)
( . .
part spore .
.
.
Acremonium, Hirsutella, Gibellula
Aschersonia . :
-
Claviceps .
Cordyceps .
Torrubiella .
Epichle .
. . )
(
.
.
Clacviceps purpurea ) ( .
.
.
.

.
Sphacelia segetum .
86
.

. .

.
8 .
Cordyceps
.
.

Elaphomyces .
Claviceps
Cordyceps militaris
.
Phyllachorales
Phyllachora, Glomerella Polystigma .

clypeus ) clypeus
( .
.
. .
.
Glomerella
80
) G. cingulata
(Colletotrichum gloeosporioides 100 .

.
. .
.
Colletotrichum
. setae .
)
(] . ) G. tucamanesis (Colletotrichum falcatum
[.
87
Polystigma Phyllachora clypeus . Polystigma

ochraceum
P. rubrum .
Phyllachora
graminis clypeus .
Diaportales

.
.
.
. .
.
Diaporthe .
.
) Phomopsis
( D. citri . D. vexans
D.
phaseolorum D. woodii . lupinosis

.
Gnomonia ) ( .
. .
Gnomonia leptostyla
) (Marssonina juglandis
.
Gaeumannomyces . .
G. graminis . .
.
Magnaporthe . .
) Magnaporthe grisea (Pyricularia grisea .
88
) (pyricularin .
.
Cryphonectria
. .
1-4 .
. Cryphonectria parasitica
)(
] Cryphonectria Endothia [.
Xylariales
.
.
.
. .
.
sympodial percurrent
. :
Diatrypaceae
.Xylariaceae
Xylariaceae .
. .
.
schizolytic

. .
) (Hypoxylon ) (Daldinia )
(Xylaria
) (Biscogniauxia ) (Camillea
.
.
- Rosellinia
89
. .
.
100 .
R. necatrix
.
Dematophora necatrix .
] . R. aquila
R. arcuata R. pepo R. quercina
R. bunodes [.
Xylaria
.
. .
X. polymorpha X. filiformis .
X. poitei .
Hypoxylon, Daldinia,
Biscogniauxia Camillea.
Diatrypaceae
Eutypa armeniacae . )
( .
Xylariaceae .
Microascales
) (
.
.
.
. Microascaceae Ceratocystidiaceae .
Ceratocystidiaceae Ceratocystis
. .
.
. Chalara .
90
blue stain sap stain .
. C. coerulescens, C. virescens
C. piliferum
C. fagacearum C. fimbriata
.
. C. fimbriata
. .
C. fagacearum .
.
Nitidulid .
.
. .
.
pressure pad .
.
. .

.
Ophiostomatales
) Europhilum (

Sporothrix Leptographium .
scolytid platypodid .
Ophiostoma epigloeum Tremella fusciformis
. . Ophiostoma
. O. ulmi .
Sporothrix Graphium
.
Hylurgopinus rufipes Scolytus multistriatus .
91
Sporothrix
. O. novo-ulmi
ceratoulmin .
Ceratocystis :
Ophiostoma

20B
Ceratocystis

21B
2B
.
-

Sordariales
.
Neurospora .
subiculum
. .
.
1-2
. .
4 Monosporascus cannonballus
. .
Sordariaceae
.
. .
1-2 .

.
.
92
Neurospora sitophila .

. Monilia sitophila .
.
.
.
8 .
.
N. tetrasperma . . Sordaria

Neurospora .
.
S. brevicolis .S. heterothallicus Gelasinospora
.
. Anixiella
.
Tripterosporaceae .
1-2 .
. Podospora
.
Podospora anserina
.
Cheatomiaceae

.
. .
.
.
Cheatomium .
Thielavia, Cheatomidium .
93
Meliolales
.

black mildew .
: mucronate
.
. capitate
. )(setae
.
.
.
2-4
) 8( .
.
Melanosporales
Melanospora .

Phialophora Gonatobotrys .
.
. fusion biotrophism .
M. zamiae
.
.
)(Discomycetses= filamentous ascomycetes with apothecia

.
94
operculate
inoperculate .
.
.
.
.
Lecanorales .
) (lignocellulosic
.
)
Ascobolous, Coprotus,
Iodophanus ( .
.
. Tuber
. :
.hymenium, hypothecium (subhymenium), excipulum

. .
excipulum .
) (ectal excipulum ) (medullary excipulum .

) (epithecium )
.(Pezizales
) (epihymenium .
:
eugymnohymenial
cleistohymenials,
.paragymnohymenial

. cleistohymenial
.
.

.
95

.
. .

. Byssonecteria . eugymnohymenial

. excipulum . paragymnohymenial
) (

.

Ostropales .
.

. rostrate dehiscense . :
-
inoperculate discomycetes
.
-
operculate discomycetes
8B
lecanoralean discomycetes .

Rhytismatales
.
. .
.
.
. Rhytisma acerinum .

.
.
96
.

.
.
.
.
.
)
( .
.

. .
.
Helotiales

.
.
.
) Leotiales (.
.
. Monilinia fructicola
Sclerotinia sclerotiorum
gladioli
Pseudopeziza trifolii
Stromatinia
Sclerotinia
cepivorum .
Sclerotiniaceae
.
.
97
. .
.
. .
.
Sclerotinia .Monilinia
.Rustromeia
:Monilinia fructicola
.
.

Monilia . ) (
. .
.
.
.
.
.
.
.
.
1-3 .

.
.
Dermataceae
excipulum
. .
. Pseudopeziza excipulum
P. trifolii . .
98
) Diplocarpon rosea (Marssonina
) D. maculatum
(Entomosporium

Pezizales
.
.
.
) (suboperculate .

. :
:Pezizaceae . .
. .
. Peziza .
.
.
.
) ( ) ( .
.
Peziza ostracoderma peat mold .
Chromellasporium ollare .
Peziza
vesiculosa
Oedocephalum .
:Tuberaceae ) (truffles .
.
. 2-4
.
99
post poral
.
.

.
. Tuber melanosporum
T. aestivum T. rufum .
EMS .
:Terfeziaceae .
.
. . Trefezia
Helianthemum
. .
- :Morchellaceae
)(pileus
) (Morchella ) (Verpa .
. ) (15-60
.
. Morchella .
. .
.
8 .
. .
.
.
Constantinella .
) M. esculenta () M. angusticeps () M. semilibera (

) M. deliciosa ( .
Verpa .

.
100
.
. Verpa bohemica 2 .
.
:Helvellaceae .
. 4 .
.
lorels .
Rhizina, Helvella,
Gyromitra Wynella Rhizina .

Helvella H. crispa .
H. undewoodi .
15 8 ) ( .
helvellic acid .
H. infula . .
Gyromitra .
.
G. esculenta G. gigas 12
. gyromitrin .
.
. G. esculenta
.
)(Loculoascomycetes
.
.
Loculoascomycetes

.
.
101
.
. .
.
.
. .

.
.
. .
. :
Elsinoe
.
. Myriangiales .
Dothidea .
.
.
. Dothideales .
Pleospora
.
. .
Pleosporales .
:
Dothideales

Dothidea .
.
6 .
- Dothideaceae
102
.
. Mycosphaerella 1000
. M. fragariae .
M. musicola M. fijiensis
.
. .
.
.
Septoria, Ramularia,
Cercospora Ovularia ] M. arachidicola Cercospora arachidicola

[.
Microthyriaceae ) .(thyrothecium

) (coroloid haustotrium .
.
Microthyrium, Trichothyrium Actinopeltis.
Myriangiales
Arthoniales .
.

.
.
. .
. :
Elsinaceae
. .
103
) (brilliant chresyl blue .
Elsine E. fowcetii . E. ampelina

E. veneta .
E. veneta
.
)
iterative germination ( . ) (Rubus
.
.
. Elsine Sphaceloma.
.
Capnodiales
subiculum
. .
.
. .
.
.
Capnodiaceae .
.
.
. .

.
Phragmocapnis .
- Antenulariellaceae
104
Scorias, Capnodium
. monilioid
. .
.
Euantenarriaceae .
. .
Pleosporales

.
.

cellular pseudoparaphyses .
.
. .
22
.
Pleosporaceae .
.
.
. :
Lewia
Pleospora
Pyrenophora
Cochliobolus
Setosphaeria
Alternaria
Stemphylium
Drechslera
Bipolaris, Curvularia
Exserohilum
Leptosphaeriaceae
. part spore
105
Phoma, Septoria Coniothyrium.
.
Leptosphaeria .
L.
maculans .
Venturiaceae .
. .
.
.
) inaequalis
(Spilocea pomi
Venturia.
V.
) V. pyrina
(Fusicladium pyrinum .
. V. inaequalis )(concentric bodies
.
Dibotryon morbosum Apiosporina Prunus
.
Apiosporina collinsii .
Amelanchier.
Hysteriaceae
.
. .

Hysterographium, Hysterium, Glonium,
Sporidesmium Septonema.
Botryosphaeriaceae .
.
Guignardia .
)
106
Discochora
Guignardia ( G. bidwellii . Vitaceae

.
Phylosticta ampelicida
.
. .
.
Melanommatales
) trabeculate pseudoparaphyses (paraphysoides
.

pseudoepithecium . .
.
. .
37B
Melanommataceae
turbinate
.
. .
. Melanomma .
subiculum .
.
38B
Arthoniales
.
.

pseuoepithecium .
semifissitunicate
. .
. .
crustose .
Roccella, Lecanactis Opegrapha.
107
Chiodecton,

Erysiphales

) (
. 10000 1617 169 44
90
. .
) ( .
.
.
.
Erysiphe polygoni
352 .
Blumeria graminis ) (
. :
Blumeria
graminis f.sp. tritici B. g. f.sp. hordei B. g. avenae B. g.

f.sp. secalis .

.
.
) (ectophytes .

.
) (hemiendophytes )
Phylactinia .(Pleochaeta Phylactinia corylea 100

.
) ( .
) (endophytes
) .(Leveillula
108
.

.
)
( .
.
.
.
)= papilla ( .
.
) (extrahaustorial matrix
.

.
.
) (green islands .
.
.
.

Cystotheca
Queirozia Blumeria Caespitotheca .

) ( .

foot cell
Blumeria
. 1-3
) 6 (
Leveillula .
(meristem
) arthrosporae basauxic
) .(acropetal )
109
Queirozia
( .
.
Leveillula, Pleochaeta
Phyllactinia . Sawadaea
.
.
) Cystotheceae
Sawadaea, Cystotheca Podosphaera(
) (fibrosin bodies
Foex . Homma
.
.
Oidium Acrosporium Oidium :
Euoidium -
. .
Pseudoidium .
Oidiopsis . .
) ( .
. Leveillula .
Ovulariopsis ) ( .
.
. Phylactinia .
Streptopodium . .
Pleochaeta .
.
) (gamocysts . ) (androgamocysts
110
) (gynogamocysts .
.
.
) (
.
. 2 8 .
) .

.
.

) (casmothecium (
.
. .
.
.
) (Cystotheca, Podosphaera 40
) Leveillula, Phyllactinia (Pleochaeta .
.
- .
Neoerysiphe .
.
.
-
) (mycelioid
Erysiphe,
.Leveillula
-
) (uncinate
.Pleochaeta, Uncinulla
111
.Phyllactinia
-
.Podosphaera
Phyllactinia
) (secretory appendage penicillate
.
Erysiphe
rosae
) Erysiphe flexuosa
( .
.

.
.
.
. Cystotheca
Brasiliomyces .

Mucor erysiphe L. Species
Plantarum . Leveill 1851
. .

.
1990 ITS 18S rDNA

ITS1-5.8-ITS2, 18S rDNA 28S
rDNA
Helotiales,
Cyttariales Rhytismatales ) Leotiomyetes ( .

) (

112
. 5
Erysipheae, Golovinomyceteae, Cystotheceae, Phyllactineae
Blumerieae
. Parauncinula Caespitotheca
.
Erysipheae
Erysiphe, Brasiliomyces Typhulochaeta .
Brasiliomyces )
( . Erysiphe
) (pseudoidium
Erysiphe
Micrsphaera,
Uncinula Uncinuliella . Erysiphe

Micosphaera, Erysiphe Uncinula
. Typhulochaeta
.
Golovinomyceteae Golovinomyces, Neoerysiphe Arthrocladiella

) euoidium ( .
Golovinomyces
Neoerysiphe Erysiphe .
.
Golovinomyces
Neoerysiphe . Arthrocladiella A. mougeotii

Golovinomyces .
Cystotheceae euoidium
. Cystotheca, Podosphaera Sawadaea .
Podosphaera Cystotheca
Sawadaea . Cystotheca
. Podosphaera
Sphaerotheca . Sawadaea
.
Phyllactineae . Leveillula
. ) (
. Phyllactinia
113
) (penicillate .
Pleochaeta
. Queirozia )(

.
Erysipheae Erysiphe Poaceae .
B. graminis euoidium .
.
Oidium phyllanthi
) Golovinomyceteae 18S (28S

. Parauncinula
. Caespitotheca
euoidium .
.
25B
>1
Oidium
Oidium
Sphaerotheca
>1
Oidium
Blumeria
>1
Oidiopsis
Leveillula
Oidium
Podosphaera
>1
Oidium
Microsphaera
>1
>1
Oidium
Ovulariopsis
Erysiphe
Uncinulla
Phyllactinia
Blumeria
Erysiphe
.
Erysiphe
) (section :
114
Erysiphe section Erysiphe - )
(pseudoidium
. 2-8 E. polygoni .
Erysiphe section Golovinomyces .
2-4 .
E.
cichoracearum.
Erysiphe section Galeopsidis . . E. galeopsidis.
2001
.
.
.
Uncinula, Microsphaera Erysiphe Sect. Erysiphe Pseudoidium Erysiphe
E. Sect. Microsphaera, E. Sect.
Uncinula E. Sect. Erysiphe .

Erysiphe Sect. Golovinomyces E. Sect. Galeopsidis Golovinomyces Neoerysiphe .
Sphaerotheca Podosphaera Podosphaera . Podosphaera ) ( P. Sect.
Sphaerotheca P. Sect. Podosphaera .
Laboulbeniales
.

) (
. :
Laboulbeniaceae,
Ceratomycetaceae, Herpomycetaceae Euceratomycetaceae

.
115
. Pyxidiophoraceae
2 .
. ) ( .

.
) (heteroecious Coelomomyces
.
Pyxidiophoraceae ) Mycorhynchidium( ) Pyxidiophora( .
.
.
. phoresy . Pyxidiophora
hypophoretic .
.

.
. )( Thaxteriola .
.
... )
3

(.
Laboulbeniaceae .
.
.
.
. ) (positional specificity
.
(sex
) of host specificity . )(direct infection

) (autoinfection
116
) (indirect infection
.
Laboulbenia
. .

.
. .

.
.
carpogonium
carpogonium
carpogenic, trichophoric trichogene

. .
Spathulosporales

Ballia Galaxaura
.
Spathulospora phycophila . Ballia
.
.
Hispidicarpomyces galaxauria Galaxaura
.
)(Basidiomycota

. :
) (mushrooms ) (toadstools ) (earth star (birds

) nest fungi ) (jelli fungi ) (bracket or shelf fungi ) (puff balls
) (stink horn fungi .
) (basidium
.
117
.
4 .
) (ballistospore .
.
.
.

.

.
.
.
Filobasidiella
) neoformans (Cryptococcus neoformans

. Agaricus
.brunnescense, Pleurotes ostreatus, Lentinula edodes
9B
.
)(somatic structures

.
) (
.
. Marasmius
. 0/5-2 4-5
. )(
) (primary ) (secondary ) (tertiary
.
.
)( .

118
.
.
.
.
.

.

) a b ( .
.

) (clamp connection
.
) ( a b
.
) (b ' b .
) (a )' (a
.
b .
.
b )
' a ' b a (b .

.
) (crozier .

)( .
.
119
Aphyllophorales
: generative, skeletal .binding

.

) (dolipore septum .

) septal pore cap (SPC
) (peranthosome . SPC
.
SPC
) (imperforate ) (perforate . SPC

.
.

.

) (basidiocarp= basidioma .
.
147 ) (Rigidioporus ulmarius .
.

.
. .
: ) (basidiole
) (cystidium
.
120
] .
) (subhymenium
. ) (trama [.
) (basidium ) 4(
.
.

.
.

4 .
) (sterigma 4
.
.
4 .
. .
:

holobasidium
phragmobasidium
.
) (teliobasidium
.

. heterobasidium
.
) Donk (1954 ) Talbot (1954
:
-
) probasidium( .
) metabasidium( .
) sterigma(
121

.
:
-
chiastobasidia
.
.
-
stichobasidia
. .
)(basidiospore

.
) (direct germination .

) (budding
. ) (indirect germination
.
.
) (Agaricales .
) ( .

(hilar
) appendix .
. ) (Buller
.
) (Wells
. McLaughlin
Boletus rubinellus
) pellicle ( .
) (ballistospore .
122
statismospore
Gasteromycetes .

.
)(asexual reproduction

) (oidia .
. )(
. .
.
.
) (oidiophore
.
) .(Coprinus lagopus
.

.
)(sexual reproduction
.
. ) (
. ) (
. )(
.
Uredinales Hymenomycetes
.
.

10 .
20-25

123
Aphyllophorales .Rhodosporidium toruloides

.

.
.
Ustilago maydis, Schizophyllum commune Coprinus
cinereus . Schizophyllum commune
) (A, A, B, B )
multiple allels( A . A B B
. A S. commune 9 A 32
B B 9 .
.

.
A
. B )
Armillaria (.
) (buller phenomenon

Coprinus cinereus
10B
) Patouillard (1900

Homobasidiomycetes
Heterobasidiomycetes .

.
Talbot (1968) .
Holobasidiomycetes Phragmobasidiomycetes, Teliomycetes .
124
)
(
Boletes .
.
Tremelloid Tremella Exidia
Auricularioid Auricularia .

. .
Hymenomycetes .
.
Gasteromycetes
.
.
. :
- 1
Ustilaginales Exobasidiales.
- 2
Uredinales .
- 3
Tremellales Hymenomycetes .
) (1996 :
Agaricales
.
) (stipe ) (pileus .
) (gills .
Boletes
. )( .
.
.
) (folicolous
) (lignicolous ) (coprophilous
. ) .(fungicolous
125

. .

14 . 6
.
.

.
) =
(fairy ring
] Marasmius oreadus .[Chlorophyllum molybdites

.

.
. ) (bioluminescense
. .
.
.
:
. . .
.
.
.
.
10 .
126
.
pH .

.

) oidia

.
( .
.

.
:
pseudoangiocarpous, hemiangiocarpous .gymnocarpous
hemiangiocarpous
.
) (inner veil .
.
.
annulus .
cortina .
Amanita ) (universal veil .

volva .

) (scales .
.
.
gymnocarpous
. pseudoangiocarpous

127
.
. ) (
) (gills
) (Boletes .
.
aequi-
hymeniferous inaequi-hymeniferous .
.
Coprinus
.
.
.
hymenophoral
trama .
.
homoiomerous
heteromerous . homoiomerous
heteromerous sphaerocysts
.
free .
:
adnate . adnexed .
decurrent .
context .
Flammulina velutipes
Coprinus
.
4 .
. ) (spore print .

.
.
. )
128
( .
. :
Agaricaceae
25
Agaricus .
volva .
. ) (
.
A. xanthodermus .
. A. campestris, A. brunnescense A. rodmani
. Lepiota Chlorophyllum
. Lepiota Amanita volva.
Boletaceae

)( )
Phylloporus ( .
.
. .
) .(Strobilomyces

Boletus . .
pulvinic acid
.
Tricholomataceae
.
Termitomyces, Flammulina, Clitocybe, Marasmius, Pleurotus Laccaria, Tricholoma
Armillaria . Termitomyces Flammulina
.
Armillaria
129
A. mellea, A. tabescens A. ostoyae .
)( .
.
.
.
A. mellea
. Pleurotus
.
.
P. ostreatus
.
.
Pseudomonas Agrobacterium
) P. ulmarius (.
Amanitaceae
Amanita
volva
annulus . . A. virosa
A. caesarea - A. muscaria
. 90-95 .
) (mycetismus A. phalloides
. 12
. volva volva .
amatoxins

phallotoxins .
Polymerase .
RNA
phalloidin
meixner
.
.
5-10
. A. muscaria
ibotenic acid(.
130
Coprinaceae

. Coprinus, Psathyrella, Panellus
. Coprinus
inaequi-hymeniferous
.
. C. comatus .
. C. cinereus
.
Russulaceae
)
(heteromerous .
.
. Russula Lactarius . Russula
. Lactarius

. L. indigo .
.
Aphyllophorales
1200 .
.
) ( :
) (pore fungi ) (club fungi ) (coral fungi ) (tooth fungi
) .(shelf and brachet fungi
) (gills .

gymnocarp .
) (unilateral
) .(amphigenous .
.
.
.
131
.
Polyporaceae, Stereaceae, Hymenochaetaceae Hericaceae
)(heart wood
.
Corticiaceae Lachnocladiaceae
.
catahymenium .
.
.

.

.
.
Aphyllophorales
Tremellales, Gasteromycetes,
Dacrymycetalse, Auriculariales Agaricales .

.
) (
.
) .(selective delignification

. )
( .
.

.
Polyporaceae .
.

132
resupinate .

. effused-reflexed
.
. context .
.
polypore . ) (trama
.
.
(Corner,
) 1932 : ) (generative hyphae

.
sclerified generative hyphae .
) hyphae :
(vegetative
) (binding hyphae
.
) (skeletal hyphae .
monomitic .
dimitic ) ( .
trimitic . :
setae KOH .
Hymenochaetaceae .
asterosetae cystidia .
.
.
cystidioles
dendrohyphidia .
.
dichohyphidia
) ( .
Lachnocladiaceae .
133
Aphyllophorales
23 .
Polyporaceae
700
.
monomitic,
dimitic trimitic . .
.
KOH
.
Fomes .
Trimitic .
conk 80 .
) (F. fascicatus, F. fomentarius .
Fomitopsis . Polyporus .
.
P.
squamosus P. sulphureus .
.
Heterobasidion annosum
.
. .
) (creosote
) (liquid borate
.
Phlebiopsis
(Peniophora) gigantea .
.

. Trametes versicolor
.
Schizophyllaceae
134
Schizophyllum commune
1-4
. .
) (split gills
.
60
.
) (mating
.
Clavariaceae

) (club fungi ) (coral fungi .
) ( .
.
.
Clavaria, Clavulina
Clavicorona.
Corticiaceae

) (resupinate .
) (lignicolous .
monomitic
. .
Laetisaria arvalis
Oomycota .
Hyphoderma
stephanocyst
.
.

. Botryobasidium
135
) (arachnoid
. Haplotrichum .
Gasteromycetes

.
) (peridium .

. )(
) (gleba . )(generative
] [monomitic ] .[dimitic
) (capilitium .
.
) (statismospore
.
. 4 .
.
.
.
) = (stinkhorns .
.
.
Nia .
. Pisolithus
tinctorius . Calvatia gigantea
9 .
5 Sclerodermatales, Lycoperdales, Phallales, Nidulariales
Tulosyomatales .
136
Lycoperdales
.
2-4 .
4-5 . Lycoperdaceae Geastraceae.
Lycoperdaceae
.
.
. .
exoperidium endoperidium .
Lycoerdon Calvatia.
Calvatia
C. gigantea .
70-80 . 7 1012
C. cyathiformis .
.
. .
.
calvatin
.
Lycoperdon .
exoperidium .
.
.
Geastraceae
) (earth stars .
. ) (exoperidium, mesoperidium

137
4-12 .
.
peristome .
Tulostomatales
) ( .

.
Calostomataceae
Calostoma
Tulostomataceae 8 . Calostoma .
Tulostoma
4-5 1 .
Battarea 30 2/5-5 .
Sclerodermatales
. 3 6
. Scleroderma
) (earth balls .
.
.
) (peridiole
.
Astraeus
.
Geastrum

) (hygroscopic movement . .
Pisolithus tinctorius )
(Pinus
. - 18
.

.
138
Phallales

.
.
. Protubera
.
. 6 Phallaceae Clathraceae
. Phallaceae
) (receptacle . Clathraceae
3-4
.
6-8
.
. Phallus
. Dictyophora indusium
.
Pseudocolus Clathraceae 3-4
. .
Mutinus .
Clathrus
.
Nidulariales
. 0/5-15
.
) (peridiole .
.

.
Cyathus stercoreus
) ( . .
Sphaerobolaceae : .Nidulariaceae
139
Nidulariaceae
6 .
.
Cyathus ) (funiculus
.
) (sheat
) (middle piece purse .

.
purse
. purse purse .

) (funicular cord . hapteron .

hapteron .
.
.
.
.
Sphaerobolus
) ( . 6
.
.
.
gemmae
. 4-8 .
Uredinales
.
. 7000
140
164 .
.

.
) (teliospore .
.
) (promycellium .
.
.

.
.

.

.
Puccinia monoica
) Arabis (Brassicaceae

) (pseudoflowers .
.
.
.
.
.
.
.
o ) (spermatia (receptive) hyphae . .
- I .
141
II . III .
IV . 5
:
) (macrocyclic 5 . ) (microcyclic .
) (demicyclic -. ) (imperfect rusts .
cataspecies .

. ) (heteroecious . o I
II III .
) (alternate hosts
o I
II III ) (primary host .
) (autoecious .

) ( .
) (endocyclic . .
- ) (Gymnosporangium juniperi-virginianae
) (Gynoconia peckiana .
] Puccinia malvacearum
.
leptoform . hemiform
.
(Puccinia
) chrysanthemi . Endophyllum
142
endoform . Puccinia punctiformis
brachyform
[.
)(spermogonium or pycnium
) (spermatia ) (receptive = flexous hyphae.
.
.

. 12 6
. .
basipetal
.
. .
.

.
spermatization
. .
.
)(aecium
.

) (disjunctor cells .
.
.
cluster cups .

. ) (peridium .
.
.
) .(peridial lip
143
.
. :
aecidium .
. Uromyces Puccinia Aecidium
.
roestelia ) ( .
.
Gymnosporangium
Roestelia .
peridermium . Cronartium, Coelosporium,
Pucciniastrum Peridermium .
caeoma
.
Melampsora, Phragmidium
Gymnoconia Caeoma .

uraecium elatraecium . uraecium

.
.
Pileolaria, Uropyxis
Ravenelia .
elateraecium Hiratsukamyces caeoma

.
) (dikaryotic haustoria .
)(uredinia
.
.
144
.
) (pedicel.
.

.
) (Melampsora )(Milesia, Pucciniastrum

) (Puccinia .
Coelosporium Chrysomyxa
aecial uredinia repeating aecia )
Gymnosporangium gaeumannii ( .

. amphispores
Hyalospora polypodii P. atrofusca P. vexans Urediniopsis
. 14
.
.
) . ( .
.
)

( .
. :
) (recognition phase .
) (signal phase . ) (parasitic phase .
pH ) .(thigmotropism
.
145
.

. ) ( .

substomatal vesicle .

. .
.
.
.
.

.
)(telium .
. .
.
.
.
Coelosporium, Cronartium, Kuhneola Chrysomyxa

leptospore
. .
) (pedicellate ) (sessile .
.
.
)(basidium .
.
4 .
.
146
.
.

.
.

. :
) (external . ) (internal

Coelosporium, Chrysospora .Ochropsora ) (semi internal

.Zaghouania
1B
2-14 .
Pucciniastraceae,
Coleosporiaceae,
Cronartiaceae,
Micronegeriaceae,
Phakopsoraceae,
Melampsoraceae, Chaconiaceae, Uropyxidiaceae, Pileolariaceae, Raveneliaceae,
Phragmidiaceae, Sphaerophragmiaceae, Pucciniaceae Pucciniosiraceae.
Pucciniaceae Melampsoraceae
. Coelosporiaceae
Cronartiaceae .

.

uredinales imperfecti .
Aecidium, Roestelia, Caeoma Peridermium Uredo

.
.
Pucciniaceae
147
.
:
Uromyces
.
. Uromyces appendiculatus U. ciceris
U. fabae U. pisi U. dianthi U. vignae
. .
Puccinia
.
.
mesospore P. heterospora Malvaceae P. .
graminis .
.
) (formae specialis=f. sp. .

Puccinia graminis f. sp. tritici . P. g. f.
sp. hordei P. g. f. sp. secalis P. g. f. sp. avenae .
:

.

.

. .
. 4
.
.
.
.
148
. 4
+ . 3 4 .
.
.

) ( .
.
.
.
.
.
. .
P. cronata P. hordei P. recondita f. sp.
tritici P. striiformis P. allii P. arachidis
P. cacabata P. carthami .
P. hordei Ornithogalum P. cronata .
Rhamnus P. recondita .
Anchusa P. triticina .
Isopyrum .
.
Puccinia .
.
Gymnosporangium
) ( .
Rosaceae Juniperaceae .
G.
juniperi-virginianae
.
.
.
149
) (telial horn
.
.
Roestelia .
.
Hemileia
.
.
H. vastatrix .

) (.
Phragmidium
.
2-3 .
Caeoma . .
P. mucronatum . P. tuberculatum
P. rubidaei .
Melampsoraceae
.
Melampsora, Cronartium, Coelosporium Masseela.
Cronartium

.
.

. .
.
.
.
150
.
4 .

.
.
.
Melampsora
.
Caeoma M. lini .
. .
Ustomycetes
.
.
Cryptobasidiales, Cryptomycocolacales, Exobasidiales, Graphiolales, Platygloeales, :
Sporidiales .Ustilaginales
Ustilaginales
.

. 1350
50 .
4000 75 .
)(
) ( .

.
.
.
.

151
. )(
. )
( .
.U. maydis
) Microbotryum violaceum (
Caryophyllaceae
. .

. )(
)
.(Ustilago striiformis
.

.
.
.
.

) (SPC
.
(clamp
) connection
.

.

) (sporidia ]
[ .
.

152
) (Ustilago scitaminea
) (Tilletia controversa .
. )
( . )(sorus
.
.

U. striiformis .

.
.
.
.
) (Ustilago maydis .
Entyloma Enthorrhizae Tilletia

) ( ) (pedicell .
.

.

. .

.

) (spore balls
.
. .
.
) (primary sporidia
153
)( .
.
. .

]
.
) Tilletia foetida
(

[.

Ustilaginaceae Tilletiaceae
. Pucciniaceae

Tilletiaceae
.
.
) (Vanky, 1985 Ustilaginaceae .

.
Graphiolaceae Graphola
Ustilaginales .
Ustillaginaceae
12B
.
.
. .

.

. . )
154
( .
.
Sphacelotheca,
13B
Ustilago Tolyposporium.
Ustilago
. U. tritici : U. nuda
U. maydis U. scitaminea U. avenae
. U. nuda U. tritici
. U. maydis a b
. a a1 a2 mating locus
. b 25 pathogenecity locus
.
bE bW
.
Tilletiaceae
.
. 2-5
8 . .

.
)
(H .
H .

H .

. :
Tilletia
T. caries . T. foetida .
.
T. controversa
6-12 T. barclayana . T. .
155
indica .
Neovossia .
Urocystis
) (spore balls
1-4 . .
U. cepulae
U. tritici .
Entorrhizae
Cyperaceae Juncaceae
.
Graphiolaceae
.
.
.
. 2-4
.
.
Graphiola Stylina .
Graphiola

Stylina
. Graphiola phoenicis
.

156

.
rDNA .

Hymenomycetes
Tremellales,
Dacrymycetales, Auriculariales, Tulasnellales Ceratobasidiales .

Sporidiales Septobasidiales
Exobasidiales
Auriculariales .
) (jelly fungi .
:
) (hypobasidium
) (epibasidium
) (sterigma .

.
Tremellales
.
:
157
-. tremelloid
-.
A 2 B -.
.
17 .
. ) (haustorial branches
Tetragoniomyces uliginosus
.
Rhizoctonia
Tremella

Tremella mesentrica .
. T. fuciformis
. .
.

.
) Wells (1994 Filobasidiaceae .
.

) (SPC .
. :
Cystofilobasidium
Filobasidium .
Filobasidiella .
.
Filobasidiella neoformans Cryptococcus neoformans
a .
.
.
basipetal .
158

. cryptococcosis

. .
Auriculariales
auricularioid .
4 .
. .
: SPC
. .
.
6 30
. .
10-15 .
. .
Phlogiotis .
.
Pseudohydnum
.
. . Tremellodendron
. Auricularia, Exidia ....
Dacrymycetales

.
) ( . .

) ( .
.
SPC .
159

) (.
.
Dacrymyces
. Calocera
.
Dacrymyces deliquescens
3 4
. .
. .
.
.
. )
( .
.
.
3-4
.
.
.
.
Ceratobasidiales
.
SPC . 6 .
Ceratobasidium Thanatephorus C. anceps .

.
Thanatephorus Ceratobasidium
T. cucumeris
Rhizoctonia
solani
.
.
160
:
.
2 1
. .
Tulasnellales
.
.
.
.
Gloeotulasnella calospora .
Rhizoctonia . 4
Tulasnella, Pseudotulasnella, Stilbotulasnella .
.
SPC Tulasnella araneosa .

Sporidiales
) Moore (1980 .
. Rhodosporidium
Rhodotorula glutinis Cryptococcoales
. Rhodosporidium sphaerocarpum :
.
. )
( .
.
.
.
)( .

rDNA .
Aessosporon salmonicolor Sporobolomyces salmonicolor
161
.
Aessosporon .
Septobasidiales
160-170 Septobasidium Uredinella .
. .

Auriculariales . .

rDNA .
Septobasidium .

. .

.

fungal garden .
. .
.
.
4
.
Uredinella .
.
.
.
Septobasidiales
Uredinales.
Exobasidiales

.
162
Theaceae,
Commelinaceae Ericaceae
.
.

.
. Exobasidiaceae 5
Exobasidium .
] Exobasidium vexans .
.

E. vaccini .
.
. [.

.
.
.
.
4-8 . .
hilum
.
. . .
Exobasidium Taphrina .
.
. .
)(Zygomycota
Zygomycetes Trichomycetes .

.
163
)(
) (zygospore .
.
. : ) coenocytic
( .

. .
-1 .
)( .
Mucorales .
-2
Entomophthorales .
-3 Zoopagales .
-4 Endogonales
.

.
.
. ) .(dimorphic

. Entomophthorales
. ) (oidium .

) (columella
) (sporangiophore .
.

.
100000 .
164
50-100
.
.
) (sporangiola ) 6-8 1-30
( .

) (merosporangium .
)
(

.
.

.
)( . ) (sugar fungi

. .
14B
9 :
Mucorales, Endogonales,
Entomophthorales, Glomales, Dimargaritales, Kickxellales, Zoopagales .

Harpellales Asellariales . Glomales
Glomeromycota .
Mucorales
) ( )( Zygomycetes .

. )
Dimargaritales .(Kickxellales
.
165
.
) (micropore
.
Mucor rouxii
. .
.
.
Choaneophora cucurbitarum
. Mucor racemosus
Rhizopus stolonifer
. Absidia corymbifer Mucor Rhizopus
Syzygites Spinellus .
)
( . sufu
tempeh .
) (rhizoid
. ) (stolon
.
.
Mycotypha Benjaminiella
. Mucor rouxii

CO2
.
.
.
. .
.
) (apophysis .
.Rhizopus stolonifer Fennellomyces
166
) (sub sporangial vesicle

.
non apophysate .Dissophora Halteromyces
Saksenaea Echinosporangium
. . Pirella
.
.
Syncephalastraceae 1-5 .
.

.
Pilobolus
.
.

.
.
.
.
. .

.Gilbertella persicaria
: .
.
.

) (zygophore .
.
.
:
167
thelemorphotic reaction zygotropic reaction -.

+ thigmotropic reaction
.

.
) (progametangia
) (fusion septum .
)(suspensor
.
)( .

) (zygosporangium
.
. ) (carbonaceous
) (non carbonaceous .

. .
. azygospore

.
. ) (opposed
) (apposed .

) (isogamous
) .(heterogamous

Absidia, Phycomyces
.Mortierella

. )
(
168
. .
. .

.
.
Mucor mucedo
) + ( .
Phycomyces nitens
)) (+( ) ( .
Rhizopus stolonifer
) (+ ) (-
) (+ )( .
1904 A.F. Blakeslee
. Burgeff 1924
. ) (
.
(trisporic
) acids .
.
. .
Rhizopus stolonifer
.

.
.
.
.
.
.
.
. .
169

.
. ) + (-
) ( .

) (prozygosporangium .
.
.
.
. 21 1-3
.
.
+ +
13 .
.
Mucoraceae
.
non
apophysate . .
.
.
. . Zygorhynchus, Rhizomucor,
Actinomucor, Parasitella, Circinomucor Mucor . Rhizomucor
.
Gilbertellaceae
Gilbertella persicaria
. .
. .
.
.
.
170
Dicranophoraceae
4 . Sporodiniella
:
Spinellus, Syzygites Dicranophora
.
Syzygites Sporodiniella .
.
Dicranophora
Spinellus 19 Syzygites Sporodiniella

. .
Absidiaceae
) (apophysate .
.
. .
Rhizopus, Absidia, Circinella
) Thermomucor ( . Absidia corymbifera .

Pilobolaceae
3 Pilobolus Pilaria Utharomyces
. .
) (

.
) (trophocyst Pilobolus Utharomyces
. ) (subsporangial vesicle
Pilobolus Utharomyces .
Pilobolus .
. Pilobolus

.
2-2/5
.
.
171
. Pilobolus hemin .
.
.
) .(apposed
. .
) (Dictyocaulus vivipores

.
Phycomycetaceae
.
. )
100000(
) .(non apophysate .
.
. Phycomyces blakesleeanus

. 1
100 .
. . -500
100 100000 .

. .
.
.
25 Phycomyces 15 .
Mortierellaceae

) (zonate . .

. .
172
.
.
. .
. Mortierella . Echinosporangium
transversalis
.
.
Saksenaceae
Saksenaea vasiformis .

.
.
) (zygomycosis .

. .
Thamnidiaceae

.
Thamnidium
. .
.
. Fenellomyces .
Cokeromyces .
Cunninghamellaceae
Cunninghamella .
.
.
173
.
.
Syncephalastraceae
Syncephalastrum
.
.
S. racemosum
S.
monosporum . .
Sigmoideomycetaceae
Sigmoideomyces Thamnocephalis
. Reticulocephalis .

.

. .
.
Thamnocephalis
Basidiobolus ranarum.
Dimargaritales
Dimargaritaceae
.
.
.
.

.
.
.
Dimargaris, Spinalia, Dispira Tieghemiomyces.
174
Kickxellales
Kickxellaceae
) (pseudophialide .

sporocladia .
Dimargaritales
(bi
) umbonate .
.
Martensella .
Kickxella
Coemansia . Dimargaritales.
Endogonales

.

Endogone
Sclerogone .
)= (sporocarps .
( . Endogone
.
.
. Endogone pisiformis
.
Endogone
pea truffles
Entomophthorales
.

. .
175

.
. .

.

.

.

hyphal bodies .
hyphal bodies .
. .

. .

.
.

capilliconidia .
.
.

capilliconidia
haptor
.
.
.
) ( 6 . 22
.
176
Entomophthoraceae
.
) 5-12( .
.
.
.
.
.
.
Entomophthora

. Entomophthora muscae (fly
) fungus .

.
.

.
.
.
2-3 .
.
. ) Strongwellsea (Erynia, Furia,
Pandora, Tarichium, Zoophthora Batkoa .
Completoriaceae
Completoria .
.
.
177
.
. .
Basidiobolaceae
.
. SPB .

9 .
Basidiobolus
ranarum .
.
.
.
. Basidiobolus

.

. .
phycomycosis
Amphoromorpha
Basidiobolus
Ancylistaceae
) 3-5( .
. .
.
.
. Conidiobolus coronatus
.
) ( .
.

.
178
.
.
Meristacraceae
) ( .
. .

. .
.
.
. Ballocephala Zygonemomyces.
Neozygitaceae
)
( .
. .
. .
.
.
) ( .
Thaxterosporium Neozygites.
Zoopagales
.
.
. .
Zoopagaceae
.
.
. .
.
. Zoophagus insidians
179
.
)
(peg .

. lethal
lollipops .
.
. .
Acaulopage
pectospora
.
Stylopage anomala .

.
.
phoresy . Zoopage Cystopage.
Cochlonemataceae
) (
.
. Cochlonema, Endocochlus Aplectosoma
Bdellospora .Amoebophilus
.
. .
.
Helicocephalidaceae
Helicocephalum, Brachymyces Rhopalomyces .
Helicocephalum
2-21

. Brachymyces Rhopalomyces
. .

180
.
.
Piptocephalidaceae
.
. )
(Piptocephalis unispora .

. Piptocephalis .

. .

.
Syncephalis Kuzuhaea .
Kuzuhaea ) (
. Syncephalis
. Piptocephalis unispora
.
.
)(Trichomycetes
.
.
.
) (
. holdfast
.
.

.
181
.
B Smittium culisetae
.
.
Smittium morbosum
.

.
.
Asellariales
Harpellales
.
Kickxellaceae .
. Harpellales ) (trichospores
.
) (exogenos

.
.
. .
Harpellales ) (biconical
.
. zygosporophore

.
.
.

.
.
4 :
182
Harpellales .
. .
Smittium Harpella Smittium .
Asellariales 2 . .
. collembolans.
Amoebidiales 2 . .
.
Amoebidium parasiticum
. Paramoebidium
.
Eccrinales . .

.
.
. .
Glomeromycota
Glomerales Glomales
. endomycorrhizae
.
.
.
.
.

) (arbuscules ) (vesicles .
183

.
.

.

.

) AM (arbuscular mycorrhizae . 80
.

.
) (
.
.
- )(
) Geosiphon pyriformis (
. .

.
AM
Glomus
.
.
AM
Endogonaceae
Endogone
Zygomycota .
Endogone
AM .
Endogone
Endogone Glomus .
184
1974 Gigaspora Acaulospora

. Endogonales
.

Glomerales 2 3 .
Glomus
2000 .

AM .

.

.
.
.
RNA
AM

Glomeromycota
.
Glomeromycetes
4 :
Archaeosporales Glomerales,
Paraglomerales .Diversisporales
Glomerales
.
.
.
Glomus .
Glomeraceae Glomus Sclerocystis .
.
Paraglomerales
) ( . .
Paraglomus Paraglomeraceae .
185
Diversisporales
.
) (auxillary cells .
Diversisporaceae,
Acaulosporaceae Gigasporaceae .
Gigasporaceae
azygospore . .
Gigaspora Scutellospora .
35B
Acaulosporaceae .
.
Acaulospora
Entrophospora .

sporiferous saccule . Acaulospora
Entrophospora .
Archaesporales

.
. Geosiphonaceae Archaeosporaceae.

] . ) (Walker, 1983
) (murograph .
.
germ shields [.
)(Chytridiomycota
Chytridiomycetes
. ) (
)
( . :
186
-.
-.
. .
100 1000 .
.

. ) (AM
. .
.
) ( .
.

.

) ( .

) (Barr, 1990
) Synchytrium endobioticum (
) Olpidium brassicae (Physoderma maydis

) () Urophylictis alfalfae ( .
Coelomomyces .
Blastocladiella emersonii Allomyces macrogynus
.
) (Neocallimastix
.
.
) (endobiotic
187
. .
) (epibiotic

] )(interbiotic
[.
) (holocarpic ) (
.

.
) (eucarpic .
.
.
)( .
. .

) (rhizomycelium .
) (monocentric
) (polycentric .
coenocytic
.
.
) (pseudoseptum
.
]
. .
) (endogenous
) monocentric
(endogenous
) (endogenous polycentric .
) (exogenous
. exogenous monocentric
188
exogenous polycentric
exogenous polycentric
colonial
filamentous . colonial sori

filamentous
[.
.
.

. .
.
) (operculum
operculate
.
inoperculate .

. :
(cell
) body ) (flagellum .
9 .9(2 ) + 2
9
3 .9(3 ) + 0 . ) (basal body or kinetosome.
. ) (fibrillar structures
.
) (rhizoplast ) (rootlet microtubules .

(dormant or
) nonfunctional kinetosome .
.

.
ribosomal aggregation .
.

189
.
) (nuclear cap .

microbody-lipid
) globule complex (MLC

.
(gamma
) particle .
.
:
-1 )(planogametic copulation
( ) = :(isogamous

.
.
( ) = :(anisogamous

.
( )( )(:
) ( .
) (
) Monoblepharidales (.
-2 )(gametangial copulation
.
-3 )(somatogamy
.
.

DNA

) .(Barr, 1992 5 Neocallimasticales,
Chytridiales, Spizellomycetales, Blastocladiales Monoblepharidale .

190
.

.
Spizellomycetales
1980 .
.
.
.
. .
Rozella, Olpidium, Caulochytrium Urophlyctis Caulochytrium .
.
C. protostelioides Cladosporium
cladosporioides .
Rozella 20
Rozella allomycis . Allomyces macrogynus
A. macrogynus .
Allomyces .
) ( .
.

.
.
Allomyces .
.
.

.
191

. .

. .
Rozella .
Rozella .
Allomyces .
host mimicry.
Olpidium 30
. .
O. brassicae
.
O. viciae . Vicia
unijuga .
.
.
.
.
.
Olpidium
Neocallimasticales
Spizellomycetales
.
. )
20( .
. MLC
rumposome 5 . 15 Neocallimastix,
Orpinomyces, Anaeromyces, Caecomyces
Piromyces .
Caecomyces Anaeromyces Orpinomyces

Piromyces Neocallimastix . Anaeromyces, Caecomyces
192
Piromyces
Orpinomyces
Neocallimastix .
Neocallimastix 9-14 .
) 1 (2
.
.
. .
. 2
.
.
.

.
) ( .
.
) (H+ H2 .
CO2 ) CH4(
.

. .
.
.
Chytridiales
.
.
.
Chytridium, Chytriomyces, Synchytrium, Rhizophydium Nowakowskiella .
.
193
.
. MLC . MLC .
Synchytrium 100 .
. exogenous polycentric
) (
sorus ) .(colonial S. endobioticum )
(
) (hypertrophy
) (hyperplasia
.
.
S. endobioticum
.
.
. .
Rhizophydium .
. R. sphaerocarpum
.
.
.

.
)
( .
Chytriomyces C. hyalinus
.
)
0/5 - ( .
.
.
.
. .
194
.
) (
. .
.

Nowakowskiella
. N. ramosa

.
. .
1/3 .
.

.
. .
.
. .
.
flexous filaments
.
.
.
.
. ) (
.
. 4-40
.
.
195
16-18
Blastocladiales

. ) (nuclear cap
.
. MLC side
body complex
. .
.
:
Catenaria
Physoderma
Coelomomyces
Allomyces, Blastocladia
Blastocladiella . Coelomomyces
) ( )
alternation of generation
(.
Allomyces .

. exogenous polycentric )(

.
.

) Emerson (1941
Allomyces
Euallomyces,
Brachyallomyces Cystogenes .
Euallomyces
. .
.
.

) .(1:1 .
196
)] (epigynous A. macrogynous
14 [ )] (hypogynous A.
arbuscula 8 [ .
.
side body .

. .
.
.
15-20 .
.
) (syrenin .
.
. ) (parisin
.
. .
.
.
.

. ) (bipolar germination

) (unipolar .
.
) (mitosporangium
) (meiosporangium .
)(
.
2-8
)( .
197
.
Blastocladiella.
Blastocladiella . B. emersonii
)
( .

. .

. .

.
.
Blastocladia )(
.
. .
B.
ramosa .
.
Coelomomyces )(
Stegomyia scutellaris
chironomids .
.
( .
.
C. psorophorae Culiseta inornata
.
) + (-
.
.
.
198

. .

.

hyphal bodies .

.
.

. Catenaria . C. anguillulae
) Coelomomyces
Coelomomyces (
.
.
.
.
.
) ( chironomids .
Physoderma
P. maydis
.

.
) ( .
.

.
turbinate cells .
199
turbinate cells .
.
Monoblepharidales

.
. Oedogoniomyces, Monoblepharella,
Gonapodya Monoblepharis .
:
.
) rumposome
(MLC Chytridiales
. Monoblepharis polymorpha

.
Monoblepharella Gonapodya .

.

. .

. .
. .

Monoblepharis .
.

.
) ( .
)( .
Monoblepharella
Monoblepharis
200
.

) ( . Gonapodya .
.
.
.
.
Oomycota
65 500-800
.
1858
Fungi
.
.

a c .
Chromista Heterokonta pseudofungi .

Heterokonta Heterokontiomycotina
Stramenopila .
:
) (tinsel ) (whiplash .
)( .
) (oospore.
.
.
) (finger print vacoules ) (dense body vacoules.

201
.
.
.
.
) 4-20 ( )
Leptomitales
Achlya Saprolegnia ( - .
) (DAP
) (AAA.
) (:
DAP
AAA
L- aspartic--semialdehyde + pyruvate - ketoglutarate + acetyl co-enzyme A
2, 3-dihydrodipicolinic acid
Homocitric acid
1
- piperidine-2, 6 dicarboxilic acid
Homoisocitric acid
N-succinyl-L--amino-- ketopimelic acid

- ketoadipic acid
N- succinyl-L- , diaminopimelic acid

L-- amino adipic acid
L,L- ,- diaminopimelic acid

L-- aminoadipic aemialdehyde
meso- ,- diaminopimelic acid

Saccharopine
L-lysine
L- lysine
) (mycolaminarins 1 3
.
(sugar alchols or acyclic
) polyoles
.
.
202
. ) (
.. ) water molds (

.

.
. ) (
Pythium insidiosum pythiosis
.

.

.
.

.
. spitzenkrper .

) (wall vesicles .
.
. ) (
) (intranuclear

) .(centric

.
)(sexual reproduction
.
.
203
) (antheridium
) (oogonium .
.
monoclinous
diclinous .
.
.
.
) (oosphere .
centrifugal centripetal
.

.

.
.
. :
exospore
epispore . endospore .
.

. .
centrifugal
Saprolegniomycetidae

. .

dense body vesicles .
. Aphanomyces
30 .
.
.
plerotic . aplerotic
204
. .
.
. .
Saprolegniales
ooplast
.
. .
:
centric 1-2
) Saprolegnia hypogyna .(S. ferax subcentric
2-3 1
) S. terrestris .(S. unispora eccentric
)
.(S. anisospora
subeccentric expersate
.
(centripetal
Peronosporomycetidae
.
205
Pythium multisporum .
.
.
.
.
apandrous.
)(asexual reproduction

.
.

. ) (auxillary zoospore
) ( )
= (principal type .
. 130 .

.
.
) (basal body . )(rootlets
1-2 .
.
.
K-bodies
. K-bodies
. MLC .
.

) (cyst . .
206
.
) ( .
: ) (mevalonate
) .(Peronosporales
.
. ) (fucosterol
Fucus . .
15B

) Oomycetes (Peronosporomycetes
).(Dick, 1995
Peronosporomycetidae SaprolegniopmycetidaeRhipidiomycetidae -
) (Barr 1983
Saprolegniopmycetidae :
-1 -2
-3
-4

-6
-5
-7
-7 .
.
4-5 . .
) (ooplasm .
centripetal . Peronosporales : .Pythiales
Peronosporales

.
.
207
. .
) ( ) ( .

. .
.
.
. .
.

.
.
.
) ( .

) (plerotic
) .(aplerotic )(
.
)( .
gonoplasm .
Peronosporaceae Albuginaceae .
Peronosporaceae ) (
. :
Peronospora tabacina
P. spinaciae .

.
.
.
. .
. :
208
Hyaloperonospora
monopodial 2

2-3 .
.Hyaloperonospora parasitica
Peronospora
.

.Peronospora rumicis
Perofascia monopodial
. .

.Perofascia lepidii
Peronospora . Basidiophora
.
Plasmopara
2-3 Bremia .
) (disk
3-5 Bremiella .

Albuginaceae
)(white rusts
.
.
. .
basipetal
. .
.
209
.
disjunctor .
)
( . .
4-12
.
. .
.
)
( .
.
. .

. Albugo :
Albugo bliti
A.
portulacae A. occidentalis A. candida )

( A. ipomoeae-panduranae .
Pythiales

) =monocentric (
. .
.
.
. .

. .
Lagenidiaceae Pythiaceae.
Lagenidiaceae
210
.
.
. Lagenidium .

.
. Olpidiopsis Lagenidium .
Olpidiopsis
Saprolegniales
. O. varians Achlya. flagellata .

.

. 24
) ( .
.
. .
.
Lagenidium
.
L. callinectes
...
.
.
. .
.
. Lagenidium 1 3 1 6
.
L. radicola
.
Pythiaceae
.
.
211
.
.
)( ) ( .
.
Phytophthora Pythium.
Pythium 120 .

.

. .
P. insidiosum
pythiosis .
.

. P. debaryanum :
.

.
. .

. .
.
.
. P. dissotocum
. .
. 7
.
. .
. .
212
. ) 28 ( ) 10-17
( .
. .
) (paragynous
. ) (hypogynous
.
Phytophthora
Phytophthora
Pythium
Phytophthora
.
50 .
)
DNA
( .
.

.
. Phytophthora infestans
1845-1846
1 1/5 .

. .
.
. 18-22
100
91 . .
.
) 24 (
) 12 ( . A1
213
A2 A1 A2 1976
.
A2 .

.
.
amphigynous .
. paragynous .
.
P. citricola .
.
Saprolegniomycetidae
.
. 3 .
.
. centrifugal.
214
Saprolegniales
Saprolegniaceae ) (water molds
.
. .
.
Aphanomyces etueiches .
Achlya, Saprolegnia Aphanomyces
. .
.

.
.
Saprolegnia

.
.
Saprolegniales Dictyuchus
.
Thraustotheca
Brevilegnia .
.

) (dimorphic
) (monomorphic .
)( .
.
) ( monoplanetic
diplanetic
. polyplanetic .
Pythiopsis . Saprolegnia
.
Achlya
215
.
adelphotaxis .
. .
Aphanomyces .

.
.
) ( . Aphanomyces
. Dictyuchus
Aphanomyces
.
.
.
) ( .
Thraustotheca
Dictyuchus
.

aplanetic .
Aplanes
Geolegnia
Achlya .

.

gemma .

.
.
:
216
. .
.
.

.

.
.
.
.
DBV .
centric, subcentric, eccentric subeccentric .
.
.
. .
.
Couch 1926 Dictyuchus Raper

Achlya bisexualis
A. ambisexualis .
monoclinous diclinous .
hypogynous .Saprolegnia hypogyna

androgynous .
Raper . Achlya
) (
) ( . Raper Achlya
. A .

Achlya . B
. C
) thigmotrophic
217
( .
A
. D
.
A B
.
Leptomitales
.
.
) (cellulin granules

. ) ( .
.
.
. .

.
)
.(Peronosporales, Rhipidiales
Leptomitus Apodachlya . .
Leptomitus lacteus
.
1-3
. .
Sclerosporales
.
. plerotic .
. Sclerospora
S. graminicola .
.
218
Rhipidiomycetidae
.
.
.
Aqualinderella
fermentans
. .
.
. centripetal
.
Aqualinderella .
.
. .
Rhipidium, Arasiospora, Sapromyces.
219

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toadstools . ) (Cladius Caesar 54

) (G. E. Clausius 1601

Rariorium Plantarum Historia .

1675 ) (van Sterbeeck

Aspergillus, Mucor, Botrytis

) Plantarum (1753 ) (Verms Chaos .

Baillard, Persoon, Link, Batasch, Schweinitz Fries

Synopsis Methodica Fungorum ) (Persoon 1801

) Systema Mycologicum (1821-1832 ) (Fries . 26

Gasteromycetes, Uredinales Ustilaginales ) (1821

6 Icones Fungorum Hucusque Cognitorum

. ) (Luis and Charles Tulasne

3 ) Selecta Fungorum Carpologia (1861-1865

1872 Grevillea . ) (Buller

) (de Bary, 1831-1888

Comparative Morphology and Biology of Fungi,

Mycetozoa and Bacteria .

Sylloge Fungorum Omnium Hacusque

Thamnidium elegans, Cladosporium

.Fusarium oxysporum, Penicillium variable pH pH

) (polykaryotic . septate hyphae .

) septal pore cap (SPC

: ) (primary septum = eu septum

) (extension zone ) (Neurospora crassa

.[myceliogenic germination ] Rhizoctonia Botrytis cinerea

mucronate hyphopodium Meliolales

] ) [spindle pole bodies (SPB SPB .

Centrosome Spindle plaque, Nucleus associated organelle

Microtubule organizing centre

pulse field gel

Leptomitus lacteus, Sclerotium rolfsii, Fusarium

Polyporus, Marasmius, Exobasidium .

Pontecorvo and Roper

) (pseudohomothallism Neurospora tetrasperma Agaricus

Trisporic acid Antheridiol Sirenin .

(Coelomomyces ) alternation of generation (.

] diplomitotic nuclear cycle

haplomitotic A nuclear cycle :

haplomitotic B nuclear cycle

)(Importance of the Fungi

ochratoxins Aspergillus ochraceous ) Penicillium viridicatum

patulin Aspergillus Penicillium )

Stachybotrys, Myrothecium, Cephalosporium Trichothecium roseum

) (lysergic acid dimethylamide = LSD -2 ) .(clavines

muscaria Clitocybe napipes

Penicillium chrysogenum 1928 .

) (kingdom Plantae ) (kingdom Animalia

) (Hogg Protoctista . 1956 ) (Copland

) (Monera 1969 ) (Whittaker

Eubacteria, Archaebacteria, Archezoa, Protozoa,

Plantae, Chromista, Animalia Fungi

Chytridiomycota, Zygomycota, Glomeromycota, Ascomycota Basidiomycota

)(International Code of Botanical Nomenclature = ICBN

Absidia corymbifer (Cohn) Saccardo & Trotter

pulchra (Sacc.) Petrak Sarcinella heterospora Sacc.

DBB (diazonium blue

- 2 ) (gametangial copulation hologamy

Dictionary of Fungi 46264

3226 32267 Alexopoulos ) (1996

Taphrinomycotina, Saccharomycotina Pezizomycotina

.Alternaria, Bipolaris, Curvularia

. basipetal .Aspergillus, Penicillium, Verticillium