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DEPARTMENT OF BOTANY,
WEST BENGAL STATE UNIVERSITY, BARASAT, KOLKATA- 700126
W.B. INDIA
ASCOMYCOTINA
(The Sac Fungi)
Gk. askos = goat skin, sac
Introduction to Ascomycotina
Sacccharomycetales
(Hemiascomycetes)
Archiascomycetes
Basidiomycetes
Classification from Alexopoulos et al. 1996
Key features of Archiascomycetes and
Hemiascomycetes that distinguish it from
Euascomycetes
Ascogenous hyphae and ascocarp are lacking in the first two
groups.
Asci are formed freely and singly either directly following
karyogamy or more rarely after a prolonged diploid phase.
Cell wall composition: very little chitin, often confined to small
ring around the site where the daughter cell is produced (the
bud scar)
Septal pore :
one or several pores may be present, usually very small and
plugged.
Hence, does not permit passage of organelles including
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Fission Yeast
Hemiascomycetes Euascomycetes
Saccharomyces cerevisiae
Features distinguishing Hemiascomycetes
from Archiascomycetes
Hemiascomycetes Archiascomycetes
Predominant growth form in May grow as a
culture as well as nature is mycelium in nature but
the yeast state although a as yeasts in the
limited mycelium or
psuedomycelium may be laboratory.
present.
Taphrina deformans
A) The assimilative mycelium is dikaryotic – different from
most other ascomycetes.
(C) The ascospores often bud, even while still inside the
ascus (left and bottom right).
(D) When the asci open to release their spores, they tend
to split across the tip, rather than around it (bottom,
left), so they are not like the rest of the operculate
group – like the asci of the Pezizales.
Members of Archiascomycetes
Ascospore budding Colony on PDA
(blastoconidium formation) a—e: Taphrina spp.
a –c Taphrina wiesneri
Fission
Taphrina and Protomyces are in the Taphrinales and are plant parasites.
Schizosaccharomyces and Saitoella are saprophytic yeasts.
Pneumocystis is a pulmonary parasite of animals.
The Saccharomycetales include brewer’s yeast and Candida albicans.
Filamentous Ascomycetes here pertains to the Euascomycetes only
Phylogeny of basal Ascomycota—nSSU and nLSU rDNA
• invagination of plasmalemma
Monokaryon
A single type of nucleus in a cell or mycelium
Dikaryon
A pair of closely associated, sexually
compatible nuclei in a cell or mycelium (n + n)
Heterokaryon (heterokaryosis)
a condition in which genetically different
nuclei are associated with the same
protoplast or the same mycelium
Parasexual Cycle
(Pontecorvo, 1956)
Establishment of heterokaryon
Mutation
Hyphal fusion
Fusion of two different nuclei to form
diploid
Haploidization by aneuploidy
From Gary Cole http://gsbs.utmb.edu/microbook/ch073.htm
INCOMPATIBILITY SYSTEMS
Homogenic incompatibility
inability of genetically similar individuals to fuse
promotes outcrossing in sexual reproduction
controlled by mating type genes (MAT)
unifactorial (bipolar)
outcrossing individuals are heterothallic
operates in gamatangia and trichogynes
Heterogenic incompatibility
inability of somatic or vegetative hyphae to fuse
somatic or vegetative incompatibility
vegetative compatibility groups (VCGs)
het genes
Mating type genes
Single genetic locus - MAT with two “alleles” not
true alleles
Idiomorphs
not homologous
encode for two distinct set of genes but at identical
chromosome location
• Heterogenic incompatibility
• Prevents the fusion of genetically different mycelia
• Multigenic and multiallelic genetic system
• het loci
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Iodine reaction of ascus walls
Application of Iodine as useful taxonomic aid to
differentiate unitunicate inoperculate type from the
bitunicate type when observed under light microscope.
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Ascus types: Prototunicate
Thin-walled ascus
Ascospores released by breakdown of wall e.g
Saccharomycetales, Eurotiales, Onygenales.
Eurotium and Gymnoascus
Ascus types: Unitunicate
Inner (endotunica or endoascus) and outer ascus walls
(exotunica or exoascus) do not separate during
ascospore release
Ascospores released through specialized adaptation at
ascus tip in the form of pore, slit or lid
Unitunicate Asci
Ascus tip
Operculate
Ascus opens by forming a lid like
operculum
Cap that detaches along preformed line
Operculum may be
lifted completely e.g Pezzizales, Sarcoscypha
coccinea or
may hinge to one side – e.g Pezzizales,
Ascobolus and Pyronema
Ascus tip
Inoperculate
No operculum
Opens by pore – e.g Heliotales like Sclerotinia or
May burst by one or two longitudinal slits at the apex
(bilabiate ascus)-e.g Pertusaria, Ascozonus.
Specialised structures found in ascus tips are generally
referred to as the apical apparatus.
Apical ring or annulus are specially
thickened inward extension of the
apical wall of the ascus arranged in
the form of cylindrical flanges. Upon
discharge of ascospore the annulus
is everted inside out like a sleeve.
Refractive ring, Neurospora
Ascus apex may be slightly thickened with single
ring Fig (a), cylindrical apex with four rings some of
which project downwards into the ascus and stains
faintly with iodine (b), pulvinus with two rings that
stains blue with iodine (c).
Annulus amyloid: when stains blue with iodine,
Amyloid ring, Xylaria
Xylaria
Dextrinoid: when stains red with Melzer’s iodine
Neurospora, Sordaria
Apices capped by a swollen plug of wall material
pierced by a narrow pore. Claviceps purpurea,
Cordyceps.
Function related to mechanism of discharge of
ascospores
Endotunica
Exotunica
Protoventuria barriae
Bitunicate (fissitunicate) Ascus
Unitunicate,
Operculate ascus
Prototunicate ascus
1. Gametangial contact
gametangia - sexual reproductive organs. Pyronema
domesticum
2. spermatization
fertilization of female gametangium by male gamete
(spermatium). Neurospora crassa
3. somatogamy
fusion of undifferentiated hyphae; rare in Ascomycota.e.g
Coprobia granulata
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Ascosporogenesis
Ascus n EMS
ascus plasmalemma
The EMS
Double membrane (dm) system. Separates from its
close association with the ascus. Plasmalemma
fragments into pieces or sheets. These dm sheets
cleave out the young ascospore
EMS
Ascus n
ascus plasmalemma
Epiplasm
Ascus n
epiplasm
ascus plasmalemma
After ascospore initials delimitation by EMS
aim
Ascus n ascospore plasmalemma
ascospore cell wall
ascus plasmalemma
In Ascomycetes meiosis occurs in the ascus similar to in basidia.
BUT two differences from Basidiomycetes:
(1) meiospores are borne inside asci and shot out;
(2) the dikaryon is a short period of the Type 1 lifecycle found
only in the ascogenous hyphal system after fertilization.
From TEXT
Ascoma or Ascocarp types
What’s inside?
Asci-scattered or in Hymenium
Centrum:- (Hamathecium + asci) excluding ascoma
wall
Hamathecium (Gr. Hama = together + theke = case):
sterile inter-ascal tissue or elements. Lacking in
Eurotiales,clavicipitales,Mycosphaerella.
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Chasmothesium
Ctenomyces Auxarthron
Apothecium
Ascocarp with asci exposed at maturity.
Cup or saucer shaped e.g Discomycetes (Pezzizales and
Helotiales). Asci unitunicate.
Sessile or without stipe, Ascobolus, Pyronema, Pezziza
With stipe, Cookenia;
Non cupulate and compound ascoma Morchella esculenta,
Helvella infula, Wynnea americana.
Epigeous: species which form their apothecia above ground.
Majority of Pezzizales.
Hypogeous: ascomata occur beneath the soil. The Truffles
Variedly coloured, fleshy and moist.
The anatomical zones
Hymenium-palisade layer of asci and paraphyses
Subhymenium-thin zone of tissue giving rise to
hymenium
Exipulum-the sterile tissue
Ectal exipulum-outermost sterile layer of apothecium
tissue of apothecium
Perithecium
Apical pore (ostiole) through with ascospores are released,
globose to flask shaped.e.g Pyrenomycetes (Sphaeriales
and Hypocreales). Asci unitunicate.
Perithecia single (Sordaria, Neurospora) or embedded in
or seated on a mass of tissue forming perithecial stroma.
Stroma brightly coloured or subued.
Ascolocular types
Loculoascomycetes = Dothideomycetes
Pseudothecia
Bitunicate asci,
Monascus locules
Asci with ascospores Bipolaris sp. (Pleosporales) Trimmatostroma abietis
Bitunicate
Broken Helicoon and Helicoma spp. (Tubeufiaceae).
ectotunica
Dothiorella sp. (Botryosphaeriales) Stroma
Trimmatostroma abietis
Conidia and
Conidia borne in pycnidium Helical conidia conidiophore Chlamydospores
Pseudothecium
Ascocarp with bitunicate asci formed in cavity
(locule) within psuedoparenchymatous
stromatic tissue also called ascostroma.
Resembles perithecium in appearance.
Venturia inaequalis
Comparision between Perithecium and
Pseudothecium
Perithecium are flask shaped Pseudothecium superficially
structure with distinct wall, the resembles a perithecium with no
peridium. distinct wall.
Without ostiole
Figs. 14-16. Schizothyrium rufulum on Encyclia sp. (Orchidaceae; Mangelsdorff 2247). 14. Longitudinal section through a
thyriothecium. Bar = 25 μm. 15. Young and mature asci embedded in ascogenous tissue. Bar = 25 μm. 16. Mature ascospores. Bar
= 10 μm.
Schizothyrium pomi
Flyspeck, on apple Without ostiole Thyriothecium cracks in
the light colored circular
area seen here
Ostiole
Figs. 7-10. Morenoina epilobii on an unknown host (Hofmann 127). 7. Surface mycelium
with young thyriothecia. Bar = 20 μm. 8. Longitudinal section through a thyriothecium with
a basal hymenium. Bar = 10 μm. 9. Young and mature asci on ascogenous tissue. Bar =
10 μm. 10. Young and mature ascospores. Bar = 5 μm.
Ascocarp Types
(a recapitulation)
Pictorial Representation of different Ascocarps
Chasmothecium
Stipitate
Apothecium
Apothecium
Compound
ascoma
Compound ascomata
Order Cyttariales Class Leotiomycetes:
Perithecium Pseudothecium
Ascostromatal types
Hysteriothecium
Pseudothecium
Thyrothecium