Division Ascomycota
Sac Fungi
I. Introduction
This is a large group of fungi, with many of significant
economic significance. Included in the group are edible forms, cheese formers,
wood rotters, producers of antibiotics and other drugs, yeasts of great
importance in baking and brewing, etc. Hyphae are septate, and cell
walls are composed mostly of chitin.
II. Asexual reproduction.
The majority of ascomycetes utilize conidiospores
or conidia.
III. Sexual reproduction.
The usual sexual spores are ascospores. These
are contained in a sac called an ascus, and are produced following
meiosis and usually a mitotic division.
Haploid mycelium of two mating types occur. These
will produce antheridia and ascogonia. Following contact
between an antheridium and ascogonium, or an ascogonium and a conidiospore
of opposite mating types, plasmogamy will occur, so that a male
nucleus will be found in the same cell as the female nucleus. This establishes
the dikaryotic state - i.e. n+n ploidy. Dikaryotic hyphae will now
be stimulated to grow from this ascogonium, in a coordinated manner with
haploid hyphae to form the fruiting body known as an ascocarp. The
dikaryotic hyphae which will actually form the asci are known as ascogenous
hyphae.
The ascus forms from the penultimate cell of an
ascogenous hypha where it forms a crozier hook. This cell undergoes
karyogamy,
followed by meiosis and usually a mitotic division, producing 8 ascospores
in an ascus. The arrangement may be ordinate or inordinate,
depending on the species. Within the ascocarp, there is usually a layer
containing the asci, and sterile cells known as paraphyses. This
layer is the hymenial layer.
The form of the ascocarp may be one of three types.
A. Apothecium - this is a more or less cup-shaped structure,
varying from quite flat to distinctly cup-like. Peziza is a good
example of this type.
B. Perithecium - this is almost closed, with a small ostiole
or opening at the tip. Ex. Claviceps. (Note - some authors include
closed ordinate ascocarps as perithecia.)
C. Cleistothecium - this is a closed ascocarp, such as Microsphaera
or Sphaerotheca. (Some authors insist that this type of ascocarp
be inordinate - see note above.)
IV. Typical Ascomycetes
A. Yeasts
These cellular organisms reproduce asexually by
budding - i.e. blastospores - and sexually by ascospores (1-8/ascus). They
show an interesting variation in the amount of life cycle spent in diploid
and haploid states. There are three genera of interest.
1. Saccharomyces cerevisiae
The familiar brewers or bakers yeast. This is one
of the oldest and most important domesticated organisms. The products of
its metabolism include alcohol, used in brewing, and CO2 which is important
in baking, causing the bread to rise. This group reproduce asexually in
both the haplophase and the diplophase. Sexual reproduction involves syngamy
and meiosis to produce 4 ascospores per ascus.
2. Schizosaccharomyces
This form is haploid, except for the brief time
following karyogamy and prior to meiosis. In this group, 8 ascospores are
formed.
3. Saccharomycodes
This for is primarily in the diplophase, and also
forms pseudohyphae. Certain cells undergo meiosis, forming 4 ascospores/ascus.
Within the ascus, two spores undergo plasmogamy and karyogamy, forming
a diploid zygote, which exits the ascus and continues as a pseudohypha.
B. Taphrinia deformans - Leaf curl fungus.
This fungus causes the crinkled leaf condition frequently
seen in peaches and other trees.
C. Powdery mildews
This group of fungi are obligate parasites, usually
occurring on the leaves of plants, where they rarely kill their hosts,
but usually slow or impair the growth of the plants. Most of them show
appendaged cleistothecia as their fruiting bodies, and produce aerial conidia
extending from the surface of the leaf. Genera include the following:
1. Erysiphe spp.
2. Sphaerotheca
3. Microsphaera
D. Endothia parasitica - Chestnut blight
From Japanese chestnut trees, where it is a hypovirulent
strain with virus-like properties. 25 years after entering the U.S., most
of the mature chestnuts were dead.
E. Ceratocystis ulmi - Dutch Elm Disease
Named by a Dutchman, but comes from Asia, where
the Chinese Elm is resistant. Carried by bark beetles, and introduced into
their passageways where the young develop. The beetle larvae use the fungus
as a food source.
F. Penicillium (= Talaromyces) - Blue mold
This genus contains species important in production
of antibiotics and in the aging of cheese. Conidia are borne on branched
conidiophores. Development of asci is a relatively recently discovered
phenomenon, and resulted in the movement from the form genus Penicillium
to the current genus Talaromyces.
1. P. notatum - the source of penicillin. Discovered by Alexander
Fleming, and developed by Florey and Chain.
2. P. camemberti and P. roquefortii - age and give the
distinctive flavors to the respective cheeses bearing their names. Originated
from caves in those regions of France, where cheeses were stored in caves
to age.
G. Aspergillus - Green mold
This genus has phialides radiating from a
head-like end to the conidiophores. It can be a serious pathogen in immune-compromised
individuals, should it enter the lungs and become systemic. It also can
produce toxic and carcinogenic byproducts (aflatoxins) if it should
grow in stored products such as peanuts. Affects humans and chickens. Generally,
however, it is a harmless saprophyte.
H. Monilinia fructicola - Brown rot fungus. Infects peaches,
plums, etc.
I. Claviceps purpurea - Ergot.
Grows on rye and other cereals. The sclerotia
contain some 20 different alkaloids, some of which cause very violent reactions
if ingested. The hallucinogen lysergic acid, vasoconstrictors, and other
substances are included. "Pit", a stimulant for childbirth is a derivative,
as are vasoconstrictors used in medicine.
Ingestion usually occurs when sclerotia are harvested
and milled with the normal grain. Violent hallucinations, gangrene and
limb loss, a burning sensation known as St. Anthony's Fire, and
death, are some of the results of ingestion. Two percent ergot in flour
will cause ergotism. Tens of thousands have died in a season. Peter the
Great's campaign against Turkey was aborted because of ergotism, and Napoleon's
defeat in Russia was partly attributable to ergotism in the ranks. There
is indication that the Salem "witches" were ergotized, thus accounting
for their unusual behavior.
The life cycle begins in the spring with the infection
of cereal flowers by ascospores. These germinate and grow into the developing
grain. The hyphae mass together and form a sclerotium which actually
replaces the grain. It is this sclerotium which may be harvested and ingested.
If it is not harvested, it will fall to the ground, and after over-wintering
perithecial
stroma or heads will grow from it. Embedded in the surface of these
stromas are many perithecia, where ascospores will eventually form. The
ascospores are long and slender, being arranged in a bundle, rather than
in a row, or a cluster.
J. Neurospora - Pink mold
This mold is very important as a genetic tool, with
its ordered array of ascospores.
K. Peziza - Cup fungus
L. Tuber - Truffles
Highly desirable underground fungi, being one of
the most expensive of mycophagists delights.
M. Morchella - morels
Another very tasty mushroom.
N. Venturia inaequalis - Apple scab
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