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Glomeromycota
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Article Contents
. Overview
. Morphogenesis (Zygomycetes)
. Sexual Reproduction (Zygomycetes)
Overview
The phylum Zygomycota consists of two classes, the
Zygomycetes and Trichomycetes. The Zygomycetes are
widely distributed and occur predominantly as saprotrophs. These fungi are readily isolated from soil, air, dung
or decaying plants. Some are parasites of other fungi,
plants or insects. Humans may experience fungal diseases
or allergic reactions to fungi. Economically important
chemicals, pharmaceuticals and foods are produced by
Zygomycetes. The other class within the Zygomycota, the
Trichomycetes, is a much smaller group whose members
occur within the guts of insects and other arthropods. The
phylum Glomeromycota contains a small number of mycorrhizal fungi that had previously been classied in the
Zygomycetes.
Molecular data conrm the importance of the Zygomycota in fungal evolution. It is likely that the ancestral fungi
diverged from the animals and plants about a billion years
ago (Bruns, 2006). These fungi were likely similar to
present-day chytrids which occur in freshwater as swimming single cells. These ancient fungi apparently lost their
agella multiple times, giving rise to nonmotile fungi that
occupied terrestrial habitats, the ancient Zygomycetes.
Even today, the Zygomycota fungi retain comparatively
primitive characteristics. Later the Zygomycete lineage
gave rise to the remaining more advanced fungi. The phylum Glomeromycota arose approximately 600 million
years ago in the Precambrian Era. The two remaining phyla
(the Ascomycota and Basidiomycota) emerged approximately 500 million years ago in the Paleozoic Era, before
the diversication of land plants (Berbee and Taylor, 2001).
Fossils of the reproductive structures of a zygomycete and
early members of the Glomeromycota have been recovered. See also: Ascomycota; Basidiomycota
A unique form of sexual reproduction characterizes
fungi in the Zygomycota. Fusion of multinucleate gametangia occurs, followed by formation of a thick-walled
zygospore. Asexual reproduction typically involves formation of nonmotile spores (sporangiospores) within a sac
(sporangium) (Benny et al., 2001; Moore-Landecker,
1996).
Morphogenesis (Zygomycetes)
Most fungi in the Zygomycota form hyphae, elongated
tubular structures. Typically, hyphae contain several haploid nuclei within a common protoplasm. Hyphae may be
divided into cellular compartments by the formation of
crosswalls (septa). Septa delimit old or damaged portions,
and reproductive structures. Septa are regularly formed
within the hyphae of some members. For many saprotrophic species, the hyphae branch extensively and rapid
growth in culture can be readily observed. See also: Hyphae
Asexual reproduction occurs after the hyphae have undergone extensive growth. Erect, aerial hyphae,
sporangiophores, arise from hyphae. The sporangiophores may be either branched or unbranched. The
sporangiophore tip enlarges to form a terminal swelling,
the sporangium. One or more sporangiospores develop
within the sporangial wall (Figure 1).
The multispored sporangium is typical for many common
and important zygomycetes such as those in the genera
Rhizopus and Mucor. In these fungi, the sporangium begins
its development as a balloon-like swelling at the
sporangiophore tip, and is later separated by a new septum.
This septum protrudes into the sporangium as a dome, the
columella. At rst, the sporangium is multinucleate. Uninucleate spores are delimited by the cleavage of the cytoplasm. They are released when the sporangial wall ruptures.
Although most of these fungi produce a large multispored sporangium, other types of sporangia occur in some
genera. Merosporangia are rodlike, containing several
sporangiospores in a row within the sporangium wall. In
others, the sporangia may contain only a few sporangiospores (these are known as sporangioles). Sporangioles are
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Figure 1 Types of sporangia formed. (a) Blakeslea trispora. Multicelled sporangium with a columella. (b) Blakeslea trispora. Sporangioles with a few
sporangiospores borne on a swollen sporangiophore tip. (c) Syncephalis. Sporangiophore bearing merosporangia. (d) Detail of merosporangium. (a, b) Gaumann
EA and Dodge CW (1928) Comparative Morphology of Fungi. New York: McGraw-Hill (after Thaxter) and (c, d) Gwynne-Vaughan HCI (1937) The Structure and
Development of the Fungi. Cambridge: Cambridge University Press (after Thaxter).
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Figure 3 Zygospores of Mucor mucedo (Mucorales). (a) Mature zygospore supported by two suspensors. (b) A germinating zygospore forming a
sporangiophore and sporangium. Bessey EA (1961) Morphology and Taxonomy of Fungi. New York: Hafner (after Brefeld).
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Class Trichomycetes
The second class, the Trichomycetes, has only approximately 200 species. These fungi produce a minute thallus
consisting of only a few cells (Lichtwardt, 1986). The
thallus is anchored to the host by a holdfast that may simply be a secretion, or a distinctive basal cell together with a
secretion. These fungi occur primarily in the midgut or
hindgut of insects and other arthropods, whereas a few
occur on the outside of the host. Distribution of the fungus
depends upon that of the hosts, and both have a cosmopolitan representation. There are two orders, the Asellariales (10 species) and Harpellales (103 species). Until
recently, the Trichomycetes was considered to have four
orders, but the orders Eccrinales and Amoebidales have
been transferred to the protozoa.
The Harpellales fungi occur in larvae of insects (such as
stoneies or mayies) that live in fresh water, especially
rapidly owing streams. The small thallus is either unbranched or branched. Branched thalli have septa that resemble those of the Kickxellales by having a central pore
and a plug. Asexual reproduction occurs by formation of
elongated monosporous sporangioles, which trail one or
more appendages after their release. The appendages become entangled in substrates at the streams bottom, where
the insect larvae often congregate and are likely to be infected by the fungus. Sexual reproduction occurs in some
species. Mating occurs between dierent thalli or between
cells of the same thallus. Enlarged cells (the gametes) fuse to
form an elongated conjugation tube within which their
protoplasts mingle. The conjugation tube gives rise to a
special branch, the zygosporophore, that bears a zygospore
at its apex. The zygospore is an elongated cell with conical
ends (Figure 4). Cytological events in the sexual cycle such as
nuclear fusion and meiosis have not been observed.
The remaining order, the Asellariales, contains only a
single family and three genera. They are widely distributed
and may occur in adult insects or isopods in streams, intertidal regions, or terrestrial habitats. These fungi produce
a branched, septate thallus that is attached to the hindgut
lining. Asexual reproduction occurs when parts of the
thallus break apart into uninucleate spores. Sexual reproduction is unknown.
anity for particular substrates. Strict soil inhabitants include species of Zygorhynchus and many species of Mucor
and Absidia. Dung-inhabiting fungi include Pilobolus and
Phycomyces. Species of Spinellus occur exclusively on decaying mushrooms. Decaying grains, fruits and vegetables
are commonly invaded by species of Rhizopus and Absidia.
Some species of Mucor can grow at 258C and have been
found in the Arctic tundra. It is not unusual to nd coldtolerant strains of Mucor or Rhizopus growing on jam or
other refrigerated food. Species of Rhizomucor are thermophilic, growing at the temperatures above 408C that
occur in insuciently dried hay, moist stored grain and
compost where temperatures rise. Some zygomycetes can
also tolerate extremes in soil pH, being adapted to either
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Figure 5 Rhizopus head rot on sunflower. Both watery soft rot and dried
shredded tissues are visible. Reproduced with permission from Harveson RH
(2007) Rhizopus head rot of sunflower in Nebraska. University of NebraskaLincoln Extension publications website.
Zygomycetes as mycoparasites
Mycoparasites are fungi that parasitize other fungi.
Among the Zygomycetes, mycoparasites occur in the Mucorales, Dimargitales and Zoopagales (Jeries and Young,
1994).
Species of Syncephalis (order Zoopagales) are necrotrophic mycoparasites of members of the Mucorales, such as
Rhizopus. Hyphae of Syncephalis invade the host hyphae,
and nally grow within the host. The hosts protoplasm is
destroyed, and eventually the host is killed. Subsequently
the necrotrophic mycoparasite can live on the dead host as
a saprotroph.
Other Zygomycetes are biotrophic mycoparasites, parasites that do not cause the death of their host fungi. The
mycoparasite contacts its host, producing an infection peg
that rst binds to the wall, and then penetrates it by mechanical and/or enzymatic means. Each infection peg develops into a specialized hyphal ending, the haustorium,
that lies between the hosts protoplast and cell wall. Haustoria absorb nutrients from the host such as vitamins
(biotin and thiamin), a growth factor (mycotrophein) and a
fatty acid (g-linolenic acid). The hyphae of the mycoparasite, as well as its reproductive structures, develop outside
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Zygomycetes as allergens
Workers in certain occupations are exposed to air that
carries an unusually large number of fungal spores, including those of saprotrophic zygomycetes. Examples include farmers who are exposed to grain, hay or silage;
woodworkers who are exposed to wood timber or pulp;
and workers in sewage disposal facilities. In these environments, the worker repeatedly inhales fungal spores that
may cause allergic reactions. Acute respiratory symptoms
such as wheezing and dry coughs, as well as fever occur.
Most cases are mild and of short duration, but in some
cases the condition may become chronic. Zygomycetes that
cause allergic reactions are Rhizopus microsporus that
grows on wood chips and timber in sawmills, and Rhizopus
stolonifer and Mucor racemosus that are encountered near
decaying vegetation or in damp interiors. Although these
zygomycetes can cause allergic reactions, they are not
among the most numerous or important of these troublesome fungi (Lacey and Crook, 1988). See also: Fungal
Spores
The disease muromycosis, caused by Mucorales members, is far more troublesome. It has been recognized in
farm animals such as pigs, horses and cattle for more than a
century. The principal fungus causing this disease in animals is Absidia corymbifera which occurs in grain and silage. Species of Rhizopus and Rhizomucor can also cause
the disease. Nodular lesions of subcutaneous tissue, lymph
nodes, or other organs such as the lungs or intestinal tract
may occur. The infection may be localized or become systemic as the fungus is spread throughout the body in the
blood, even infecting the brain. Some animals show no
symptoms, whereas others develop pneumonia. Animals
having a systemic infection can have dysfunctions of the
digestive system (anorexia and persistent diarrhoea) or
neurological disturbances if the brain is involved. Mucormycosis is a common cause of abortion in cattle and
pigs.
Species of Rhizopus (especially Rhizopus oryzae) and
Mucor cause muromycosis in humans. Infection of humans
is not common and occurs primarily in individuals with a
serious existing condition such as diabetes, leukaemia, organ transplantation or malnutrition. In patients with diabetes, mucormycosis originates in the sinus and nose.
Early symptoms include fever and dull sinus pain, followed
in a few days by double vision and increasing fever. Fungal
invasion of the ophthalmic artery leads to blindness and a
coma can result when the fungus invades the frontal lobe of
the brain. Without proper treatment, the diabetic patient
may die within a few days or weeks. Progression of muromycosis may be dierent for other patients, with the infection beginning in the lungs or gastrointestinal tract. In
all cases, there is the likelihood that the fungus will spread
rapidly through the blood to the brain, causing death
(Schipper and Stalpers, 2003). See also: Immunity to
Fungi; Infections in the Immunocompromised Host
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Organic compounds
These fungi produce various useful enzymes and metabolites (Table 1).
Commercial production of proteases and lactic acid by
these fungi is especially signicant.
Rhizomucor miehei and Rhizomucor pusillus (along with
one other fungus) produce acid proteases that are used to
manufacture about half of the cheeses worldwide (Lowe,
1992). Cheese manufacturers had previously relied upon
suckling calves as a source of rennin (chymosin), a milkclotting enzyme produced in the fourth stomach. A worldwide shortage of calves rennin occurred because cheese
consumption increased while calves are usually slaughtered
at a later age. This shortage resulted in a search for an
alternative milk-clotting agent, and it was found that fungal proteases could substitute for calves rennin. Fungal
proteases can clot the milk, but the activity is not so intense
that the protein is completely hydrolysed. This is a desirable feature that is not shared by proteases from bacteria,
mature cows or pigs. Further, many vegetarians seek
cheese that was not manufactured with enzymes from animals. Fungal proteases have other uses (Table 1).
Rhizopus oryzae can produce signicant amounts of lactic acid from glucose.A high yielding strain originated as a
mutant, and further successive selection of six mutants led
to a strain of Rhizopus oryzae that can convert up to 90%
of glucose to lactic acid, representing a substantial
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Producer
Use of product
Enzymes
Amylase
Rhizopus oryzae
Galactosidase
Lipases
Pectinase
Proteases
Metabolites
b-carotene
Fumaric acid
Lactic acid
g-linolenic acid
Triglycerides
Mortierella vinaceae
Rhizomucor miehei
Rhizomucor pusillus
Rhizopus oryzae
Rhizopus oryzae
Rhizomucor miehi
Rhizomucor pusillus
Blakeslea trispora
Phycomyces blakesleeanus
Rhizopus delemar
Rhizopus oryzae
Mortierella spp.
Mucor spp.
Mortierella vinaceae
Steroid bioconversion
Steroids occur in humans as hormones produced by the
testes, ovaries, placenta and the adrenal cortex. They can be
useful as anti-inammatory agents, for the treatment of
sexual disorders or regulation of fertility.
Steroids may be obtained from natural sources such as
ox bile, urine or plants, or they may be synthesized. They
dier from each other in their side-chains, and their eectiveness as a drug depends upon a particular structure.
Chemists sometimes modify the steroid nucleus or the steroids isolated from natural sources. However, transformations or additions of side-chains may be dicult to
accomplish by chemical means, or may be too expensive
or time-consuming. Some bacteria, actinomycetes and
fungi can modify the steroid precursor if it is added to the
culture medium. An example is the hydroxylation of carbon-11, which can be accomplished by Rhizopus stolonifer
but is dicult for the chemist to accomplish. The particular
conversion is required to produce cortisone from progesterone. Species of Rhizopus and Mucor can also accomplish
other conversions such as hydroxylation of other carbons,
side-chain cleavage, epoxidation and hydrogenation. In the
commercial production of specic steroids, the steroid nucleus is often converted into the nal form by a combination of chemical procedures and conversion by specic
microbes.
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Phylum Glomeromycota
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transport these minerals into the root. The uptake of phosphorous and other minerals enhances plant growth, particularly in phosphorous-decient soils. VAM roots also
have a greater capacity to take up water, and are better able
to withstand drought conditions than plants lacking VAM
fungi. The ability of these fungi to increase mineral and
water uptake is related to the increased absorptive surface.
Very narrow hyphae grow into smaller spaces in the soil
where the larger roots and root hairs cannot penetrate, as
well as to grow away from the root and into new zones
where minerals are available. Hyphae may grow from one
plant to another, forming a nutritional network. Later,
some species form terminal swellings on hyphae within the
root. These are vesicles, which are similar to spores as they
have thick walls and a high lipid content. Vesicles may serve
as storage organs or survival units under adverse conditions (Linderman, 1997). See also: Mycorrhiza
References
Further Reading
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