Life cycles, spore formation and germination

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L.2 –Biotechnology
Mycology
D.Ebtihal Muiz
Life cycles, spore formation and germination
The times at which a fungus reproduces sexually and asexually, the
nuclear condition and morphology of these stages are known as the life
cycle of that fungus. The life cycles of a fungus reflects the adaptations of
that organism to its environment and/or niche, and is of great interest
from an evolutionary and biological point of view. A complete
understanding of an organisms life cycle is necessary if we want to
control or use a particular species.
As you might expect for the fungi, life cycles are not as consistent as
those found for flowering plants. The life cycle of an organism can be
described as the series of events from zygote to zygote. Superimposed on
the life cycle is often an asexual reproductive phase - a manner of
increasing numbers of individuals without going through sexual
reproduction (fusion, karyogamy and meiosis).
Asexual reproduction depends on mitotic division of the nucleus,
requires less expenditure of energy and reduces the role of chance. On the
negative side, it provides less genetic variation and relies on mutation and
the parasexual cycle as a source of variation. The structures formed
during asexual reproduction include:
zoospores (Chytridiomycota), sporangiospores (Zygomycota) and
conidia (Ascomycetes, Basidiomycetes). The generic term for asexual
spores is mitospore.
Sexual reproduction requires the fusing of two cells and/or nuclei,
sometimes through the production of specialized structures such as
gametangia. Thus the process involves chance (since the two mates must
find each other) and is energetically more expensive than asexual
reproduction On the positive side it provides new combinations of alleles.
Structures formed during meiosis include: resting sporangia giving rise
to zoospores (Chytridiomycota); zygospores (Zygomycota); asci and
ascospores (Ascomycota); basidia and basidiospores (Basidiomycetes).
One thing is quite clear in the fungi, their life cycles are quite
different from those of higher plants and animals where the organism is
diploid and meiosis occurs in cells which are totally dependent on the
diploid organisms to produce haploid cells which are short-lived since
they must fuse to form a zygote or die.
Haploid-diploid - these phases alternate regularly, an unusual
condition in fungi restricted almost entirely to a few species of aquatic
Chyridiomycota.
Diploid - the haploid phase is restricted to the gametes or
gametangial phase. Most of the Oomycota may conform to this pattern
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and it is more similar to your life cycle and that of flowering plants. This
supports the alignment of this group with the Stramenopila.
Spore Germination
Spores are generally considered dormant structures. Dormancy is
any rest period or reversible interruption of the phenotypic development
of the organism. The spore stage is a quiescent phase situated in the life
cycle of a fungus between two phases of active growth. There are two
basic types of dormancy: constitutional (memnospores) and exogenous
(xenospores). Constitutional dormancy is due to an innate property of the
species. For example a spore might have a physical or chemical barrier in
the wall that inhibits the uptake of water or nutrients or there may be a
metabolic block that needs to be overcome or an inhibitor that must be
leached out before germination will occur. With exogenous dormancy,
theoretically the spore is capable of germinating immediately after it is
produced but it remains dormant due to unfavorable environmental
conditions such as water availability, temperature, pH, etc.
Factors required for germination.
I. Water. Water is essential for spore germination and the amount
required varies among taxa.
80% humidity - Aspergillus, Penicillium
90-95% - Alternaria, colletotrichum, Ustilago, Cladosporium
More that 95% - Venturia, Magnesia
Liquid water - many species including Endoconidia, peronosporales,
Sclerotinia conidia, rust spores, etc.
2. Temperature. The temperature required for germination varies with
the species and often reflects geographical distribution, habitat and life
cycle.
3. Nutrition. Many spores carry with them internally all the nutrients
needed but some require a special exogenous factor, e.g., vitamins,
inorganic ions, particular carbon source.
4. pH. Spores of most fungal species will germinate at a pH between 4.56.5.
5. Oxygen. Small quantities of oxygen are needed for spore germination
and low quantities of oxygen are not thought to limit germination.
Changes during spore germination.
1. Water content increases and the spore swells due to the absorption of
water. The
spore comes out of the quiescent metabolic state as enzymes go into
solution.
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2. Vacuolation increases.
3. Endoplasmic reticulum and other cell organelles increase.
4. Respiration increases and the mitochondria enlarge and form more
cristae.
5. Vesicles thought to be involved in wall synthesis also appear.
6. In spores with lipid bodies, the lipid bodies disappear as the lipids are
used up as an
energy source.
Spores in general:
by wet weight spores generally contain 25% protein and 20% fat. and
they have a low water content relative to vegetative mycelium. Cell walls
of spores are generally not fibrillar, but
they are multi—layered and often contain melanin and have
Ornamentations.
Spores contain all normal mycelial organelles. Respirators reserves
include lipids, glycogen, phospholipids and polysaccharides that can
include sugar alcohols Like Trehalose), Respiration rates in spores are
only 1-4% those of vegetative mycelium, but obviously the more reserves
a spore has, the longer it will survive.
Dormancy:
Dormancy occurs when spores do not immediately germinate after
formation. Dormancy is a break in the life cycle. There are two types,
endogenous (constitutive) and exogenous (induced). Endogenous
dormancy is clue to sonic internal quality of the spore, a barrier to water
or nutrient entry, a metabolic block, or an inhibitor. Self inhibition
prevents spores from germinating in dense suspensions. It can be by
excessive sensitivity to oxygen or carbon dioxide levels, nutrient
competition, or most usually due to the presence of inhibitors. These
molecules are often active in the 1-10 nanomolar range. These inhibitors
have to be leached away before germination takes place.
There can also lie physical barriers to germination. In one of the athlete
foot fungi, Microsporium gypseum, there is a protein layer around the
spore which prevents the uptake of water. This layer is removed by the
action of a fungal acid phosphatase enzyme. This enzyme is inhibited by
high levels of phosphate, and until phosphate levels in the environment
drop the fungus spore does not germinate.
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Endogenous (induced) dormancy occurs because of some external
condition, and whilst these conditions prevail the spore will not geminate.
As soon as the limiting factor is removed the spore germinates.
Table I . A summary of the characteristics of fungal spores with
endogenous and exogenous dormancy:
Endogenous dormancy:
Exogenous dormancy:
Displaced from point
of origin Remain at porn of origin
Definite launch mechanisms
Released by autolysis
Small and thin walled spores
Large and thick walled
Short survival time
Survive for a long time
Germinate readily tinder suitable
Germinate after a specific
conditions
stimulus or removal of an inhibitor
Optimal environmental signals trigger the end of dormancy and the
onset of germination. Chemical stimuli can trigger germination. This is
frequently seen in pathogens where host compounds can act as
germination stimulants.
Germination begins with imbibition, the uptake of water, which can cause
a 3 to 20 fold increase in size. Spherical growth also accounts for some of
the swelling. Eventually polarized growth starts, with the emergence of a
germ tube from the spore. The spore wall may he ruptured and a new cell
wall covered germ tube emerges, or the spore wall may be softened and
the germ tube then emerges.
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General View of fungi
What is a fungus?
1. Heterotroic organism devoid of chlorophyll
2. Has cell walls
3. Non motile (exception is zoospores)
4. Reproduces by spores
5. Primary storage product is glycogen
6. Usually filamentous, eukaryotic and multicellular
Hypha (pl. hyphae)
1. Filaments constituting the body (thallus) of a fungus.
2. Hyphae elongate by apical grow th.
3. Hyphac arc referred to, collectively, as mycelium (pl. mycelia).
4. Hyphae have cross walls called septa (sing. septum).
5. Hyphac with septa are known as septate.
6. Hyphae without septa arc known as aseptate or coenocytic.
7. Fungal cell wall consists of B linked glucans and chitin.
Hyphae organized into structures
1. Stroma (pl. Stromata)— compact, somatic structures that is cushion
-like on or in which fruiting bodies are formea.
2. Sclerotium (pl, sclerotia) — hard testing body resistant to unfavorable
conditions.
3. Rhizomrph —strands to thick shoestring—like Structures that exhibit a
high degree of internal structure where hyphae may lose their
individuality and have a thick rind and growing tip.
Hyphal behavior may indicate type of organism
1.Nectroph (Perthotroph) — uses enzymes or toxins to kill host cells in
advance of their hypltae and then hyphae grow between dead or dying
cells.
2. Biotroph (obligate parasite) — obtains nutrients only from living cells,
They have specialized hyphal branches (haustorium —pl. haustoria) that
penetrate the host cell wall and invaginate the host cell membrane
3. Hemibiotroph — requires living host cells but soon act like nectrophs
(Colitotrichium, lindemuthianumn).
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Other hyphal structures
1. Appressorium (pl. appressoria) — specialized infection structures that
form at the tips of hyphae.
2. Appressoria adhere to the host surface and form penetration pegs.
3. Penetration pegs can penetrate hard artificial surfaces such as gold foil
and nonbiodegradable Mylar membranes.
4. Protoappressoria — slight hyphal tip swellings that act like an
appressoria.
5. Reproduction in Fungi
6. Fungi have both asexual (anamorphic) and sexual (teleomorphic)
reproduction.
7. Some fungi are holocarpic (entire fungus converts into one or more
reproductive structures).
8. Most fungi are eucarpic (reproductive organs arise from only a portion
of the fungus body — thallus — pl. thalli).
Asexual Reproduction
1. Fragmentation of somatic hyphae or throcpores (thallic conidia)
2. Fission of somatic cells into daughter cells
3. Budding
4. Mitotic spores
Asexual spores
I. Chlamydospores arc thick walled thaIlic conidia that function as a
resting spore
2. Asexual spores are either sporangiospores or conidia.
3. Sporangiospores vs. Conidia
4. Sporangiospores are borne in a sac called a sporangium. The cytoplasm
within the sac is cleaved into sporangiospoies. The sporangium is borne
on a
specialized hypha known as a sporangiophore.
5. A conidium is borne on the tip or side of a specialized hypha known as
a conidiophore.
Sexual reproduction in Fungi
1. Plasmogamy — (p!) fusion of protoplasts (bring nuclei close together).
N+N
2. Karyogamy (k!) — fusion of nuclei. 2N
3. Meiosis (mi!)— reduction of chromosome number to haploid state. N
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4. In some fungi, k! follows p! quickly; in others, K! is separated from p!
by time and space.
Sex Spores in Fungi
1. Most plant pathogenic fungi exist as N or N+N and are only 2N in the
zygote.
2. In the Oomycota, individuals are 2N for most of their life cycle.
3. Some fungi have hyphae with different genotypes of nuclei within the
same hyphal cells. This is known as heterokaryosis. Such individuals are
referred to as being heterokaryotic.
4. Gametes in Fungi
5. Sexual organs — gametangia
6. Isogamete I isogametangia — male and female are indistinguishable
7. heterogametes I heterogametangia — male and female are
morphologically different
8. Male — antheridium; femaIe — oogonium
9. Classification
How Fungi Obtain Nutrients
All fungi obtain food by secreting digestive enzymes and then absorbing
the organic molecules produced (external digestion).
— extensive hyphae network provides enormous surface area for
absorption
— many fungi able to break down cellulose in wood
Ecology of Fungi
• Fungi and bacteria are the principal decomposers in the biosphere.
— mineral cycling
Fungi are virtually the only organisms capable of breaking down lignin.
• Fungi often act as disease—causing organisms for both plants arid
animals.
—agricultural damage
— human health
• Mutualistic associations
— lichens — fungi and green algae
— mycorrhizae — fungi and plant roots
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