Ascomycetes

Introduction

The phylum Ascomycota

(also called ascomycetes)
largest group of
In most ascomycetes the ascus
fungi, more than 32 000 species
contains
in 3400 genera
eight ascospores
and turgid
The name is derived from the
ejecting its spores
Greek words
by
askos = a leather bottle, bag or bladder
a squirt mechanism
Mykes = a fungus
so ascomycetes are
SAC FUNGI

The
characteristic feature of the group is
that the
sexually produced spores, the ascospores
are contained within a sac, the
ascus.
HABITAT

There is a very wide range of The range of habitats is wide, as would

lifestyles be expected of such a large and diverse
Some ascomycetes group of
are fungi
saprotrophs Ascomycetes grow in soil,
necrotrophic are common on
or the above-ground parts of plants
biotrophic parasites of plants found in freshwater & sea
and animals, including humans.

Many ascomycetes Most ascomycetes are recognized by

grow as endophytes in their
symptomless associations fruit bodies or ascocarps, i.e. the
with plants. structures
which surround the asci.
Some are
mutualistic symbionts, for
example the lichens
Vegetative structure

Ascomycetes may grow either as yeasts,

i.e.
unicells
multiplying by budding or fission,
or
as mycelia consisting of septate hyphae

Some fungi may switch from the

yeast to the filamentous state or vice
versa,
i.e. they are DIMORPHIC

A good example of a
dimorphic fungus is Candida
Vegetative structure

The mycelial septa of ascomycetes

are usually This means that
incomplete organelles
developing as such as
mitochondria & nuclei
transverse centripetal
are
flange-like ingrowths from the free to travel
cylindrical wall of from cell to cell
a hypha
which fail to meet at the centre the large
nuclei are constricted as
so that they pass through the
in most pore
ascomycete septa there is
one central pore permitting
cytoplasmic continuity and
streaming between adjacent
segments of mycelium
Vegetative structure They measure 150-500nm in width
and are sufficiently large to block the
Individual cells may be septal pore.
uni- or multinucleate They rapidly do so near regions where
& Usually one
the cytoplasmic continuity Woronin body blocks one pore.
between the cells means that
the mycelium
of an ascomycete
is effectively coenocytic
Proteinaceous organelles termed
WORONIN
bodies may be closely grouped
near the central pore

Woronin bodies
are globose structures or
‘hexagonal’ (polyhedral)
crystals made up essentially of
one protein
and surrounded by a unit
membrane.
Vegetative structure

The blockage in
the septal pore is consolidated by
deposition of
further material

Woronin bodies are formed near the

hyphal apex
&
are
transported to
more distal regions of the hypha as
septa develop.
Vegetative structure

HETEROKARYOTIC mycelia
The mycelium of many also occur and generally
ascomycetes is arise through anastomosis, i.e. the
HOMOKARYOTIC cytoplasmic fusion of vegetative hyphae.
(Gr. homoios = like, resembling;
karyon = a nut, meaning Following anastomosis (cross connection)
nucleus), i.e. all nuclei between homokaryons of differing
in a given mycelium are genotypes, nuclei, other organelles and
genetically identical. plasmids may be transferred between
one mycelium and
another so that a given mycelium or even
a
single cell may contain nuclei of
different kinds.
Vegetative structure

However, the ability to form

heterokaryons is
under genetic control and a degree of
genetic
similarity between homokaryons is
necessary
for it to occur.

Failure to establish a heterokaryon

is a phenomenon known as
Heterokaryon incompatibility
or
vegetative incompatibility
Vegetative structure Because the different types of
nuclei do not always divide at the same
Mycelial anastomosis between rate,
homokaryons, the ratio of nuclear types in a
nuclear division succeeded by heterokaryotic
migration may mycelium may change with time and
result in the rapid spread of an respond to
introduced changes in external conditions such as
nucleus into a mycelium, thus nutrient availability.
transforming a
homokaryon into a This gives the mycelium a degree of
heterokaryon. genetic flexibility which sometimes
An ascomycete manifests
mycelium may thus consist of a itself as the formation of sectors in a
mosaic of cells, mycelium
some of which are in agar culture
homokaryotic and others Another important
heterokaryotic. source of genetic variability which may
arise within a heterokaryon is the
parasexual cycle, a
process in which genetic recombination is
brought about in the absence of meiosis.
Life cycle
Ascomycetes reproduce by
[Link] reproduction
[Link] reproduction
The mycelium is often
capable of asexual reproduction, e.g. by
Sexual reproduction
fragmentation,
Sexual life cycles in the budding or
strict sense, i.e. involving by the formation of conidia,
nuclear fusion and meiosis, chlamydospores,
occur only in those sclerotia, etc.
ascomycetes which possess
asci, because it is
within the young ascus that
these events occur.
Ascospores of most
ascomycetes contain one or
more haploid nuclei, and
therefore most
(but by no means all)
ascomycetes have a haploid
vegetative mycelium.
Life cycle

The structure and

formation of conidia is described
below.
Some
yeasts,
e.g. Saccharomyces cerevisiae
show an alternation
of diploid and haploid yeast-like
states
and here the diploid state is the
commonly
encountered form
in contrast to
Schizosaccharomyces in
which the
vegetative cells are haploid
Life cycle
The mating behaviour of
ascomycetes may
be homothallic or heterothallic.

In HOMOTHALLIC In HETEROTHALLIC ASCOMYCETES

ascomycetes the mycelium the ascus usually contains four
derived from a single ascospores
ascospore is capable of reproducing of one mating type
sexually, and
i.e. by developing asci. four of the other.
Sexual reproduction

Plasmogamy is of three main types:

[Link]-gametangiogamy

Fusion occurs
between differentiated gametangia.

An example
is Pyronema domesticum
where fusion
is between
the Trichogyne, a filamentous
extension of the large, swollen ‘female’
gametangium (the ascogonium)
and a
less swollen ‘male’ gametangium, the
antheridium,
which donates nuclei to the trichogyne
and thereby to the ascogonium
Sexual reproduction

2. Gameto-gametangiogamy.

Fusion takes place

between a small unicellular male gamete
(spermatium) and a differentiated
female
gametangium (ascogonium).

The spermatium
is rarely capable of independent
germination
and growth and may only germinate to
produce a short conjugation tube which
fuses with the wall of the ascogonium.
An
example is Neurospora crassa in which
the
spermatium fuses with a Trichogyne
Sexual reproduction

3. Somatogamy

Fusion takes place

between
undifferentiated hyphae
i.e. there are no recognizable sexual organs

This type of
sexual behaviour is shown by
Coprobia granulata
whose orange ascocarps are common on
cattle dung.
Asexual life cycles

Asexual (imperfect) stage accomplished

by
Sporulation (formation of spore)
Vegetative methods

[Link]: the conidia are exogenously

produced, non motile, deciduous asexual
spores.
Conidiospores are produced on
reproductive hyphae called conidiophore

It may be branched or unbranched

In Penicillium the conidiophores are long

& usually branched
Ultimate branch bears a cluster of
bottle-shaped structure called
Sterigmata each of which produces a
long chain of conidia
Asexual life cycles
2. Oidia 3. Chlamydospore

Some Ascomycetes the hyphae don’t Large, thick-walled spore like resting
form spores in succession cells
Which contain reserve food & resistant
at their free ends. To drying & remain dormant for long
periods.
Instead septate hyphae undergo
additional septation and break up Chlamydospore may be terminal
Or fragement under proper
Conditions into their component cells. Under favorable condition
Chlamydospores germinate
Single celled fragments behave as &
asexual spores called oidia Each producing a new mycelium
Asexual life cycles

1. Fragmentation
Separation from the mycelium
of hyphae or segment 2. Fission
of hyphae called the fragments. Ascomycetes with a unicellular thallus
Fragment as
consist of fission yeasts usually multiply by this
one or more living cells, method.
Mother cell elongates.
repeated cell division Nucleus divide into two.
& apical growth, The daughter nuclei move apart.
each fragment Ring-like ingrowth of the wall material
produces appears at the wall in the middle of the
a new mycelium cell.
It grows inwards covered in front by the
plasma membrane.
It meets in the centre stretching across
the cell forming a complete partition
called the septum.
The septum thickens & then splits
into two layers,
one for each daughter cell before they
separate.
Asexual life cycles
3. Budding
Saccharomyces reproduce by budding

Cell protoplast covered by cell wall

Bulges out in the form of
Small protuberance at or near one
Pole of yeast cell.

Protuberance gradually increase in

Size & called bud.
Daughter nuclei migrates into enlarging
Bud.
Nucleus of parent cell along with
The vacuole divides
It grows & becomes constricted
At the base.

Opening between the bud & yeast cell

closes,
double wall forms
& two cells become physiologically
distinct.
They separate from each other
3. Parasexual reproduction

Process in which genetic recombination

can occur through
nuclear fusion & crossing over
of chromosomes during mitosis.

Meiosis does not occur

&
instead haploidization
takes place
by the
successive loss of chromosomes
during mitotic divisions.
It is believed that the
necessary cytological steps take place in
a regular
sequence which Pontecorvo (1956) has
termed
the parasexual cycle.
3. Parasexual reproduction

The essential steps include

(i) nuclear fusion between genetically distinct
haploid nuclei in a heterokaryon to form
diploid nuclei
(ii) multiplication of the diploid nuclei along with the original
haploid nuclei
(iii) the development of a diploid homokaryon
(iv) genetic recombination by crossing-over during mitosis in
some of the diploid nuclei
(v) haploidization of some of the diploid
nuclei by progressive loss of chromosomes (aneuploidy)
during mitosis
3. Parasexual reproduction

This process was discovered in

Emericella
Aspergillus nidulans
which can reproduce sexually
by forming asci
Genetic mapping based on gene
&
recombination
asexually by forming
following conventional sexual
conidia
reproduction
has been compared with mapping
By changing the nutrient
based on
content of the medium on which the
parasexual recombination and has
fungus is grown,
yielded identical
the development of asci and therefore
results.
of
normal sexual reproduction can be
prevented.
Economic Importance of Ascomycetes

1. Neurospora crassa has been the subject of intensive genetical

research related to its relatively simple nutrient requirements,
rapid growth, its capacity to produce mutants and the ease with which
it can be grown and cross-mated in culture.
2. The dissection of ascospores from its asci by micromanipulation has
enabled tetrad analysis to be performed.
3. Research on this fungus led to the important one-gene-one-enzyme
concept.
4. Budding yeast (Saccharomyces cerevisiae) and a
fission yeast (Schizosaccharomyces pombe) were amongst the
first eukaryotes for which the entire genome was sequenced.
5. Studies on S. cerevisiae were basic to the understanding of the
biochemistry of anaerobic respiration whilst studies of S. pombe
have provided key facts by which to interpret the fundamental process
of cell division, which in turn has a bearing on the understanding of the
apparently uncontrolled growth of cancerous cells.
Economic Importance of Ascomycetes

The economic significance of fermentation processes involving

ascomycetes and their conidial relatives is immense

Alcoholic fermentations by yeasts as the basis of the

wine and brewing industries,
antibacterial antibiotics
such as penicillin from Penicillium chrysogenum
and cephalosporin from Acremonium spp.,
and organic acids such as
citric acid from Aspergillus niger
The immunosuppressant drug
cyclosporin, which reduces the tissue rejection
response and thus facilitates organ transplants,
is a metabolite of Tolypocladium inflatum
Economic Importance of Ascomycetes

Ascomycetes are important in food production as

in bread-making by yeast,
cheese ripening by
Penicillium roqueforti & P. camemberti &
the fermentation of soybeans and wheat
by Aspergillus,
yeasts and bacteria to produce soy sauce.
The mycoprotein Quorn is produced
from mycelial biomass of Fusarium venenatum

direct use of ascocarps as food or

food flavourings are morels (Morchella spp.) and
truffles (Tuber spp.).
Economic Importance of Ascomycetes

However, food spoilage may result from ascomycete contamination.

A well-known example
is contamination of cereal grains and grass
by sclerotia of the ergot fungus Claviceps purpurea, which can cause
severe, sometimes fatal, neurological, muscular and circulatory
diseases such as
gangrene or abortion in cattle
and man.

Studies on the alkaloid toxins

contained in ergot sclerotia led to the discovery
of drugs useful in obstetrics and the treatment
of migraine, and in the identification of the hallucinogen lysergic acid
Economic Importance of Ascomycetes
Another potentially
serious mycotoxin is aflatoxin produced in
groundnuts, cereals and other foodstuffs
infected by Aspergillus flavus.

Aflatoxins are
highly carcinogenic in poultry and mammals,
including man.

Other mycotoxins include

Zearalenone from Gibberella zeae, which causes
infertility in cattle and pigs

Trichothecenes from Trichothecium roseum and Fusarium spp.,

which cause aleukia in farm animals and
man.
Economic Importance of Ascomycetes

A family of plant growth hormones,

the gibberellins, now produced commercially,
were discovered in an investigation of Bakanae
(foolish seedling) disease of rice.

It is not surprising that such a large group

as the Ascomycota should contain numerous
pathogens of plants and animals.

Lifestyles
are similarly varied, including biotrophic, hemibiotrophic
and necrotrophic associations.

Many ascomycote pathogens are of considerable economic

importance.