Zygomycota

Secondary article
Article Contents

Elizabeth Moore-Landecker, Rowan University, Glassboro, New Jersey, USA

. Overview

The division Zygomycota is an important and diverse group of fungi, containing two
classes, the Zygomycetes and Trichomycetes.

. Morphogenesis
. Sexual Reproduction
. Biodiversity

Overview

. Ecology

The class Zygomycetes contains almost 900 known species,

which 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 animals. The other class within the Zygomycota,
the Trichomycetes, is a much smaller group (fewer than 200
known species) whose members occur within the guts of
insects and other arthropods. There, these small fungi
absorb nutrients from the guts contents without harming
the arthropod.
A study of the 18S ribosomal RNA of the Zygomycetes
and other fungi helps conrm the importance of the
Zygomycetes in fungal evolution. The ancestral fungi
occupied aquatic habitats, and had spores that swam about
by beating their agella. The Zygomycetes evolved from
these ancient fungi about 500 million years ago, at the
beginning of the Silurian era. One of their major
characteristics was the loss of agella and the move to
terrestrial habitats. Later, about 400 million years ago, the
Zygomycetes gave rise to the most advanced fungi, those
belonging to the Ascomycota and the Basidiomycota
(Berbee and Taylor, 1993). Fossil representatives of the
Zygomycetes have been found in remains from the
Carboniferous era (Pirozynski, 1976).
A unique form of sexual reproduction characterizes
fungi in the Zygomycota. Sexual reproduction involves
formation of a resting spore, the zygospore, that results
from the fusion of gametangia. Asexual reproduction
typically involves the formation of nonmotile spores (the
sporangiospores) within a sac (the sporangium).

Morphogenesis
The majority of fungi in the Zygomycota form hyphae and
a mycelium. The role of the hyphae is to absorb nutrients
from the substrate. Typically, the hyphae are nonseptate
and contain several haploid nuclei. Later septa form to
delimit old or damaged portions, and to delimit the various
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.

. Economic Importance of the Mucorales

(Zygomycetes)

Asexual reproduction occurs after the mycelium has

undergone extensive growth. Erect, aerial hyphae, sporangiophores, arise from the mycelium. The sporangiophores may be either branched or unbranched. The tips of
the sporangiophores enlarge 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 tip of the sporangiophore. A new septum
is formed at the base of the sporangium, delimiting it from
the sporangiophore. This septum protrudes into the
sporangium as a dome, and is known as a 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, and these are known as sporangioles. Sporangioles
are much smaller than the multispored sporangia and lack
a columella. In some cases, only a single sporangiospore is
formed. Monosporous sporangioles are deciduous, and
the sporangiospores and the sporangium are disseminated
as a single unit (some mycologists refer to a deciduous
sporangium as a conidium).
Asexual reproduction can occur by chlamydospores in
some species. Chlamydospores are formed as the hyphae
undergoes some dierentiation, forming cells that have
thick walls and dense cytoplasm.
Upon germination, the sporangiospores or chlamydospores form a new mycelium.

Sexual Reproduction
Most species in the order Mucorales share a common,
characteristic pattern of development. With minor varia-

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Zygomycota

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); c, d, Gwynne-Vaughan HCI (1937) The
Structure and Development of the Fungi Cambridge: Cambridge University Press (after Thaxter).)

tions, this pattern exemplies sexual reproduction in other

Zygomycetes.
Depending upon the species, sexual reproduction may or
may not require two genetically dierent strains of
mycelium. Those fungi that can reproduce without a
second genetic strain are said to be homothallic, while
those that require two dierent, compatible strains for
reproduction are heterothallic. Heterothallism in the
Mucorales is controlled by two alleles at a single genetic
locus controlling incompatibility. The mating types are
designated A and a, or ( 1 ) and ( 2 ). A compatible cross
can occur only between two dierent mating types (A and
a, but not A and A or a and a).
In a heterothallic fungus such as Mucor or Rhizopus, the
hyphae of the ( 1 ) mating type grow towards the hyphae of
the ( 2 ) mating type. Attraction of the hyphae towards
each other may occur as a result of volatile hormones. The
plus strains produce 4-hydroxymethyltrisporates, while
the minus strains produce trisporins. These volatiles induce
2

progametangia formation, the enlarged clubshaped ends

of the hyphae. Each progametangium is subsequently
separated by a septum into the terminal multinucleated
gametangium, and the subterminal suspensor. In most of
the species, the suspensor and gametangia are equal in size.
Two gametangia (one from each mating type) come into
contact with each other and adhere tightly together. The
wall breaks down between the gametangia, allowing the
cytoplasm and their component nuclei to mingle to form a
single cell, the zygote. At rst, the zygote is surrounded by
the thin wall derived from the fused gametangia. The
zygote begins its development into the zygospore as a new
thick wall forms within the thin outer wall, which is
eventually sloughed o (Figure 2). The mature zygospores
are typically dark (black or brown) in colour, and have
thick walls roughened with warts or other ornamentations.
By the time the zygospore has undergone a resting period
and has begun to germinate, some nuclei will have fused
with each other to form diploid nuclei, which subsequently

Zygomycota

undergo meiosis. Typically, some nuclei disintegrate. As

germination occurs, a hypha or a sporangiophore terminated by a sporangium emerges from the zygospore
(Figure 3).
The above description is characteristic for most species
of the Mucorales. Variations, however, do occur. In
various genera, the gametangia and suspensor may be
unequal in size (Zygorhynchus). The progametangia may
become curved and tonglike in appearance (Phycomyces)
(Figure 4). The zygosporangium may arise as a budlike
outgrowth from the larger of two fused gametangia
(Endogone). Sexual hyphae may remain undierentiated
before zygospore formation in some species.

Under some conditions, such as poor cultural conditions, a gametangium may fail to fuse with a second
gametangium. Such a gametangium sometimes develops
into an azygospore, a resting spore that resembles the
zygospore but is smaller. Unlike the zygospore, the
azygospore is not involved in nuclear events that characterize the sexual cycle. Some fungi routinely produce
azygospores but not zygospores.

Biodiversity
Class Zygomycetes
The larger class, the Zygomycetes, contains almost 900
known species. These fungi are commonly collected
worldwide. Within this class, important dierences in
asexual reproduction and ecological adaptations occur
among the orders. Orders in the class Zygomycetes are:
Dimargaritales (14 species)
Endogonales (21 species)
Entomophthorales (185 species)

Figure 2 Stages in the development of a zygospore of Sporodinia grandis

shown in a median view. Nuclei are not shown. (a) Two swollen hyphal tips
come into contact. (b) A septum separates the gametangia from the
suspensors. The gametangia then fuse, and the intervening walls break
down, allowing the cytoplasm to mingle. (c) The zygote continues to
enlarge. (d) A thick wall develops surrounding the protoplast of the zygote.
This wall lies to the inside of the original walls of the zygote seen in the
earlier stage. (Gaumann EA and Dodge CW (1928) Comparative
Morphology of Fungi New York: McGraw-Hill (after Keene).)

Figure 3 Zygospores of Mucor mucedo, a member of the 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).)

Zygomycota

Figure 5 Basidiobolus ranarum, a member of the Entomophthorales.

Unequal gametangia (left) are involved in the formation of the zygospore
(right).  990. (Gwynne-Vaughan HCI (1937) The Structure and
Development of the Fungi Cambridge: Cambridge University Press (after
Fairchild).)

Figure 4 Pilobolus kleinii, a member of the Mucorales. (a) Sporangiophore

bearing sporangium. The sporangium will be forcibly discharged into
space. (b) Zygospore borne by unequal suspensors. (Bessey EA (1961)
Morphology and Taxonomy of Fungi New York: Hafner (after Buller).)

Glomales (160 species)

Kickxellales (21 species)
Mucorales (122 species)
Zoopagales (161 species)
The largest of the above orders is the Mucorales (3
families, 122 species). The majority are saprotrophs. Wellknown and important genera include Mucor, Rhizopus and
Absidia. The mycelium is nonseptate, except in old
portions and at the base of reproductive structures.
Asexual reproduction most often involves multispored
sporangia, but can also involve few- to one-spored
sporangioles or merosporangia. Species of Pilobolus can
eject the entire sporangium out into the open and away
from the dung heap where they live. The phototropic
sporangiophore bends towards the light (an opening in the
heap of dung), internal osmotic pressure builds up within
the sporangiophore which suddenly bursts, forcibly carrying the sporangium away on a minute jet stream of uid.
The sporangium can land on fresh grass where it can be
consumed by a herbivore.
The Kickxellales is represented by only one family, the
Kickxellaceae, which also has a cosmopolitan distribution
with most organisms living as saprotrophs. These fungi
reproduce asexually by forming monosporous sporangioles. Young as well as old mycelia are divided by septa,
each septum having a central plug. The smooth zygospores
are formed from hyphae that are not dierentiated as
progametangia. The small order Dimargitales bears many
similarities to the Kickxellales, but diers primarily by
producing merosporangia containing two spores. These
4

fungi are obligate mycoparasites of members of the

Mucorales.
Fungi in the Entomophthorales are almost all specialized as parasites of insects or other animals. Some are
saprotrophs in soil or dung. The mycelium consists of both
long and short segments. The shorter segments are the
thick-walled hyphal bodies. Asexual reproduction can
occur when the mycelium breaks up into the hyphal bodies,
or when these hyphal bodies bud. Also monosporous
sporangioles are formed that are often forcibly discharged.
Once discharged, a sporangiospore can germinate to form
a secondary sporangiospore. Sexual reproduction occurs
by the fusion of similar or dissimilar gametangia to form a
zygospore having a thick two- or three-layered wall. The
zygospores may be hyaline or dark, and smooth or
ornamented (Figure 5).
The majority of the fungi in the Zoopagales live on other
fungi, nematodes, amoeba and small terrestrial animals.
Predacious species can trap living protozoa such as
amoeba, or even living nematodes. These fungi form a
nonseptate mycelium that produces a sticky secretion that
adheres to the living prey. Subsequently, the host is
invaded by ne, branched haustoria and its entire body
contents are consumed by the fungus. Those species of the
Zoopagales that are endoparasitic typically produce a
coiled thallus within their host. After death of the host, the
fungus reproduces asexually by forming monosporous
sporangioles, and also reproduces sexually by forming
spherical zygospores, covered by hemispherical warts
(Figure 6).
The Endogonales is represented by a single genus,
Endogone. These fungi are saprotrophs in soil, or are
externally associated with roots. All of the fungi in the
Glomales are associated with roots where they form
mycorrhizae. These fungi are widespread in the temperate
zone. They can form multicelled and macroscopic sporocarps up to 25 mm in diameter. The sporocarps occur
chiey beneath the surface of the ground but can occur
above ground in litter. Sexual reproduction is known only
in some members of Endogone, which produce sporocarps

Zygomycota

Figure 6 Cochlonema verrucosum, a member of the Zoopagales that killed an amoeba. (a) A coiled thallus is present inside the dead amoeba and is
producing two chains of monosporous sporangiospores (conidia). (b) Zygospores forming outside of the remains of the amoeba. (From Drechsler C (1935)
Mycologia 27: 640.)

containing zygospores. Some members of the Glomales

(such as species of Glomus and Sclerocystis) produce
sporocarps that contain chlamydospores. Such sporocarps
may occur in addition to loose clusters of chlamydospores
that occur in the soil. Some fungi lack sporocarps entirely,
for example Acaulospora laevis (Glomales), which produces azygospores loose within the soil. The zygospores,
azygospores and chlamydospores are all thick-walled
spores that undergo a resting period.

Class Trichomycetes
The second class, the Trichomycetes, represents only about
200 known species. Unlike the members of the Zygomycetes, these fungi do not produce an extensive mycelium,
but instead produce a minute thallus (body) consisting of
only a few cells.
The small, inconspicuous thallus is anchored to the host
by a holdfast. The holdfast may simply be a secretion, or
may consist of a morphologically distinct basal cell
together with a secretion. The branched thalli have septa
that resemble those of the Kickxellales by having a central
pore and a plug. The unbranched thalli are coenocytic.

These fungi occur primarily in the guts of insects and other

arthropods, while a few occur on the outside of the host.
The majority of the hosts occur in freshwater habitats,
while some occur in marine waters or are terrestrial. The
four orders assigned to the Trichomycetes are:
Amoebidiales (10 species)
Asellariales (10 species)
Eccrinales (62 species)
Harpellales (103 species)
The Harpellales contains the largest number of species
(at least 103), which occur in the larvae of insects living in
fresh water (especially rapidly owing streams). These
fungi form a small thallus that may be either unbranched or
branched. Asexual reproduction occurs by the formation
of elongated monosporous sporangioles, which trail one or
more appendages after their release. Sexual reproduction
occurs in some species. Mating may occur between
dierent thalli or between cells of the same thallus. As
the mating process is initiated, the uninucleate cells
(gametes) enlarge, contact each other, and join to form
an elongated conjugation tube. The walls separating the
gametes quickly dissolve, allowing the cytoplasm to
intermingle. The conjugation tube gives rise to a special
branch, the zygosporophore, which bears the zygospore at
5

Zygomycota

Figure 8 Enterobryus elegans, a trichomycete in the order Eccrinales.

Young thalli are attached to the gut of the host. These thalli have not yet
become septate to form sporangia (Bessey EA (1961) Morphology and
Taxonomy of Fungi New York: Hafner (after Leidy).)

reproduction occurs. Members of the Asellariales produce

a branched and septate thallus, and reproduce asexually
when parts of the thallus break apart into uninucleate
arthrospores, which then fall apart from each other. The
coenocytic thallus produced by members of the Eccrinales
is unbranched, or only branched at the base (Figure 8). The
thallus cleaves into sporangia, beginning at the apex and
proceeding downward. Each sporangium forms a single
sporangiospore, which then escapes.

Ecology
Zygomycetes as saprotrophs

Figure 7 A zygospore of a trichomycete borne on a zygosporophore.

(From Moss ST and Lichtwardt RW (1977) Canadian Journal of Botany 55:
30993110).

its apex. The zygospore is an elongated cell with conical

ends (Figure 7). Cytological events in the sexual cycle such
as nuclear fusion and meiosis have not been observed.
The remaining three orders share many similarities with
the Harpellales, but may dier in their thallus structure and
mode of asexual reproduction. Sexual reproduction is
almost universally absent in these orders. Members of the
Amoebidiales form an unbranched, nonseptate thallus
that converts entirely to sporangiospores when asexual
6

Most members of the Mucorales and Kickxellales are

saprotrophs, feeding and living upon dead organic matter.
These zygomycetes are readily isolated from the soil, dung,
decaying plant debris and old mushrooms. There is some
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. Stored grain and decaying
fruits and vegetables are commonly invaded by species of
Rhizopus and Absidia, resulting in considerable loss due to
their activities. Species of Rhizomucor are thermophilic,
growing at the temperatures above 408C that occur in
closely packed vegetation, grain and compost where
temperatures rise.

Zygomycota

Growth of saprotrophic fungi on organic debris is of

extreme importance in nature because it provides a means
to break down natures garbage, returning the elements to
the soil and atmosphere in a form that can be used by other
organisms. The Zygomycetes are among the rst fungi to
colonize a fresh substrate such as plant remains or dung
that has just reached the ground. The spores germinate
rapidly, and their hyphae grow rapidly on the fresh
substrate, giving them an advantage over competing fungi.
For a short time, the Zygomycetes are the dominant fungi
on the substrate. They utilize the readily available sugars
and other nutrients. Later, their numbers decline because
they are unable to tolerate their own byproducts of
metabolism (especially carbon dioxide from respiration)
and the competition from other fungi. Along with some
other soil fungi, species of Rhizopus and Mucor can
detoxify some pesticides in soil, rendering them harmless.

Zygomycetes as allergens
Workers in certain occupations are exposed to air that
carries an unusually large number of fungal spores.
Examples include farmers who are exposed to grain or
silage, and other individuals who work indoors in foodprocessing establishments, harvest mushrooms, or work
with timber or wood pulp. In these environments, the
worker is repeatedly exposed to the fungal spores which are
inhaled and 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. Although
the Zygomycetes are not among the most numerous or
important of these troublesome fungi, a variety of
Rhizopus microsporus that grows on wood chips and
timber in sawmills has been identied as an allergen for
Swedish woodworkers (Lacey and Crook, 1988).

Zygomycetes as pathogens of mammals

Zygomycetes as mycoparasites
Mycoparasites are fungi that parasitize other fungi.
Among the Zygomycetes, mycoparasites occur in the
Mucorales, Dimargitales, and Zoopagales.
Species of Syncephalis (order Zoopagales) are mycoparasites of members of the Mucorales, such as Rhizopus.
Hyphae of Syncephalis recognize and invade the host
hyphae, and nally grow within the host. The hosts
protoplasm is destroyed, and eventually the host is killed.
Such a mycoparasite that kills its host is known as a
necrotrophic mycoparasite. 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 protoplast and the wall of the host
hypha. Haustoria absorb nutrients from the host such as
vitamins (biotin and thiamin), a growth factor (mycotrophein), and a fatty acid (gamma-linolenic acid). The
hyphae of the mycoparasite, as well as its reproductive
structures, develop outside of the host. The physiology of
biotrophic parasites is nely tuned to that of the host so
that the host is not killed. This is advantageous to the
mycoparasite because it ensures a continued source of
nutrients for these fungi, which would be unable to live as
saprotrophs on the dead hosts. Obligate biotrophic
mycoparasites include fungi in the genera Dimargaris and
Dispira (order Dimargitales) and Piptocephalis virginiana
(order Zoopagales). These fungi are almost exclusively
restricted to host fungi occurring in the Mucorales.

As noted above, the majority of the Zygomycetes live

primarily as saprotrophs in the soil, on decaying plant
material, and on dung of various animals. Some species are
thermotolerant and grow on stored grain and silage, where
temperatures become high. Consequently, the sporangiospores of these fungi are present in the air breathed by
mammals. The fungi rarely cause a problem, but are
capable of becoming pathogenic to individuals who are in a
weakened condition from factors such as malnutrition,
immunosuppression or diseases such as diabetes. Zygomycetes are known to cause diseases of various vertebrates
including a variety of farm animals as well as humans.
Pathogenic Zygomycetes occur in the orders Entomophthorales and Mucorales.
Pathogenic fungi in the Entomophthorales primarily
aect the subcutaneous tissue and nasal passages of horses
and humans. The symptoms include subcutaneous granulomas that become ulcerated, or nodular growths of the
nasal passages that can cause mechanical blockage. The
infection may spread to the lymph nodes. The fungi
responsible are Entomophthora coronata, which is also an
insect parasite, and Basidiobolus ranarum, which normally
occurs in the gut of frogs. In the temperate zone, these fungi
particularly infect horses. Infection of humans is known
primarily from Indonesia. A disease caused by either of
these fungi is known as entomophthoromycosis.
The disease termed mucormycosis caused by members of
the Mucorales is far more troublesome, and 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, a thermotolerant
fungus that can be readily isolated from grain and silage.
Certain species of Rhizopus and Rhizomucor can also cause
the disease. Nodular lesions of the subcutaneous tissue, the
lymph nodes, or other organs such as the lungs or intestinal
7

Zygomycota

tract may occur. The infection may be localized or may

become systemic as the fungus is widely disseminated to
various parts of the body through the blood. Even the brain
may become infected by the fungus. Some animals show no
symptoms of the disease, while 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 R.oryzae) and Mucor are
the principal pathogens causing mucormycosis in humans.
Infection of humans is not common and occurs primarily
in individuals with a serious preexisting condition such as
diabetes, leukaemia, organ transplantation or malnutrition. In patients with diabetes, mucormycosis originates in
the sinus and nose. Early symptoms include a low grade
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 mucormycosis
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.

Zygomycetes as insect parasites

Species of Entomophthora and Massospora (order Entomophthorales) are specialized parasites of insects.
Entomophthora infections begin when a sporangium
comes into contact with the host. Germination of the
sporangium occurs and the resulting germ tube penetrates
the thin-walled integument occurring between segments or
joints of appendages, and enters the body cavity. Within
the host, the fungus mycelium converts into a number of
short, thick hyphal bodies that grow by budding and
ssion. Eventually these hyphal bodies replace much of the
internal structures of the host, resulting in its death. Upon
death, the host is anchored to its nal resting place by
specialized hyphae. Sporangiophores emerge through the
intersegmental areas, and each sporangiophore produces a
single monosporous sporangiole, which is forcibly discharged the following day. Zygospores develop within the
body cavity. A familiar example of such an infection is
caused by Entomophthora muscae, which kills houseies,
frequently observed on window panes surrounded by a
ring of discharged sporangia.
Entomogenous fungi may help control populations
of insect pests in nature. The gypsy moth can defoliate
forest trees and can become a serious problem. The
populations of the gypsy moth have been held in check
in the northeastern USA because this insect is parasitized
8

by Entomophaga maimaiga (order Entomopthorales),

which can sometimes occur in epidemic proportions.

Zygomycetes as mycorrhizal symbionts

Mycorrhizae are symbiotic associations between the roots
of vascular plants and certain fungi. The symbiosis is
mutualistic, providing selective advantages to both the
fungal and plant partners.
The vesicular-arbuscular mycocorrhizae (VAM) fungi
represent a special subclass of the mycorrhizal fungi. The
VAM fungi are Zygomycetes belonging to the order
Glomales, and in particular, species of Glomus. The
mycorrhizal relationship is obligate for these fungi, and
in nature, they are found only in association with a root.
They form chlamydospores or azygospores, but not
zygospores. The spores are usually produced singly, but
can be formed in sporocarps. Dispersal of the spores can
occur by wind, water or animals such as worms or insects.
Spores in hypogeous sporocarps are typically dispersed by
a mammal that digs up and eats the sporocarps. After
dispersal, the spores germinate and the fungi grow as
hyphal strands or wefts over the roots. Some of the hyphae
extend into the soil, while other hyphae invade root hairs,
the epidermis and perhaps the cortex as well. The fungus
produces enlarged globose vesicles within some root cells,
and dichotomously branched hyphal endings, the arbuscules, in other root cells. The vesicles apparently serve as
storage organs or survival units under adverse conditions.
Nutrient transfer between the root cell and fungus
probably occurs primarily across the arbuscules, which
have a relatively large amount of surface area in contact
with the cytoplasm of the root cell because of their
branching. Senescent arbuscules can also undergo lysis,
which also makes nutrients available to the host. The VAM
fungi depend entirely upon the plant for soluble carbohydrates, which they are unable to obtain as saprotrophs.
Having the VAM fungi can favour the plant, increasing its
growth. The fungi increase the transport of minerals
(especially phosphates) into the plant which enhances
plant growth in poor soils. VA roots also have a greater
capacity to take up water, and are better able to withstand
drought conditions than plants lacking the mycorrhizal
fungi. The ability of these fungi to increase mineral and
water uptake is in part related to their ability to increase the
absorptive surface by having very narrow hyphae that
grow into the smaller spaces 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.

Trichomycetes as commensals
The Trichomycetes may be found when the gut of an
arthropod host (such as an insect, millipede, or crab) is
dissected. Within the gut, the thalli are attached to the gut

Zygomycota

lining and lie within the gut lumen. The thallus is rmly
anchored to the gut lining by a holdfast, but does not
penetrate the gut tissue. Only the largest of these fungi can
be observed without a microscope. When clustered
together, the longer thalli may make the gut appear to be
fuzzy or hairy, giving the name Trichomycetes (hair
fungi) to this group of fungi. In a few species, the thalli may
protrude from the anus of the host and can therefore be
detected without dissection. One trichomycete grows
externally on the exoskeleton of aquatic arthropods.
The hosts are insects and other arthropods. The majority
of the Trichyomycetes occur in freshwater habitats on
hosts such as the larvae of ies and other insects. Millipedes
from terrestrial habitats are particularly likely to be hosts,
but Trichomycetes also occur on some other terrestrial
arthropods such as beetles and isopods. Marine animals
that are likely to be hosts are various crabs that live in the
intertidal or tidal zones. The distribution of both the hosts
and the Trichomycetes is worldwide.
The Trichomycetes must live within a suitable host. The
host typically shows no response, either favourable or
unfavourable, to the presence of the fungi. The fungi,
however, assimilate nutrients from the materials within the
hosts gut. The relationship is considered to be commensal
in nature. Only a few Trichomycete species have been
isolated and grown in pure culture, and therefore our
knowledge of their nutritional requirements and relationship with the host is incomplete. Although the relationship
is obligate for the fungi, it is not obligate for the hosts.
Continuation of the life cycle depends upon the eective
dispersal of the fungal spores (sporangiospores and
zygospores) produced within the hosts gut. The spores
are shed into the environment with the faeces, when the gut
lining is shed as moulting occurs, or when the host dies. The
arthropod hosts all have chewing mouthparts and feed
upon debris or algae. As they feed, they consume the fungal
spores, which then are stimulated to germinate within the
gut to produce new thalli.

Economic Importance of the Mucorales

(Zygomycetes)
Some species of the Mucorales are of economic importance. Rhizopus and Mucor have been used in the
commercial production of lactic acid, fumaric acid and
succinic acid. Blakeslea trispora and Phycomyces blakeleeanus synthesize carotene pigment, which can be used as a
colouring agent for margarine and baked goods. The
enzymes amylase and amyloglucosidase, produced by
species of Rhizopus, are useful in the hydrolysis of starch
during beer production or for the removal of starch
sizing from fabrics. Mucor pusillus is a source of protease,
which is used to tenderize meat, and also in cheese and
bread production. Especially important uses of these fungi

are in the bioconversion of steroids and Oriental food

production.

Steroid bioconversion
Steroids occur in the human as cholesterol, an essential
component of cell membranes; and as hormones produced
by the testes, ovaries, placenta and the adrenal cortex.
Steroids can be useful as anti-inammatory agents, for the
treatment of sexual disorders of regulation of fertility.
Steroids may be obtained from a natural source such as
ox bile, urine or a plant, or they may be synthesized in the
laboratory. The steroids dier from each other in their
side-chains, and their eectiveness as a drug depends upon
a particular structure. Chemists can sometimes readily
modify the steroid nucleus or the steroids isolated from
natural sources. In some cases, however, transformations
or additions of side-chains may be very 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 present in the
culture medium. An example is the hydroxylation of
carbon-11, which can be accomplished in culture 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 important conversions such as hydroxylation of other carbons, side-chain
cleavage, epoxidation and hydrogenation. For the microbial modication, the steroid is dissolved in a solvent and
added to the culture medium in which the fungus is
growing. Aerobic conditions are maintained during the 24
to 48 h incubation period; during this time the microbe can
modify the steroid that is subsequently chemically extracted from the medium. 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.

Oriental foods
Various foods are prepared by culturing the raw products
together with appropriate microbes. Familiar examples
include the conversion of milk to yogurt by bacteria, and
the processing of milk curd into Roquefort cheese by a
fungus. Foods that are prepared in such a manner have
probably been valued in various cultures before the
beginning of recorded history. The texture, avour and
nutritional properties of foods are often improved by
microbial growth, while spoilage is delayed. In oriental
countries, diverse foods are prepared from soya beans,
wheat and rice through the deliberate or accidental
inoculation with various microbes, including members of
the Mucorales. Two oriental foods prepared with fungi
belonging to the Mucorales are tempeh and sufu.
9

Zygomycota

Tempeh is a solid food prepared from soya beans in

Indonesia. The soya beans are cooked, drained, cooled,
and then innoculated with spores of Rhizopus oligosporus.
The inoculum is transferred in a small amount of tempeh
made previously, or from a wrapper that had held a cake of
tempeh. Once inoculated, the soya beans are wrapped in a
large leaf (such as a banana leaf) or placed in a perforated
plastic bag. The fungus is allowed to grow on the soya
beans for about 24 h before being consumed. The tempeh
can be fried, roasted, or added to soups. Tempeh is more
readily digested than the soya beans from which it is
prepared, and the nutritive value is increased as well. The
fungus actively secretes abundant quantities of proteolytic
enzymes that convert proteins into amino acids, and
lipolytic enzymes that break down fats. Riboavin, niacin
and vitamin B12 each increase in quantity.
The Chinese prepare sufu, which resembles a creamy
cheese. Soyabeans are ground and the milk is pressed from
the mash. The soya bean milk is precipitated with vinegar
or calcium sulfate to produce a solid curd which is then
pressed into a mould such as wooden box to remove excess
water and to form a cake, now known as tofu. Tofu can be
used as a food, or processed further into sufu. For sufu
production, cubes of tofu are inoculated with Actinomucor
elegans or a species of Mucor (Mucorales) and are
incubated for 3 to 7 days. During this time, the fungus
produces a solid mycelial lm over the surface of the soya
bean curd. Meanwhile the fungus produces both proteolytic and lipolytic enzymes, as occurs in the production of
tempeh. The mouldy cubes are then immersed in a brine
containing salt and rice wine and allowed to age for about 2
months. The cheese and brine are bottled and sterilized for
marketing.

References
Berbee ML and Taylor JW (1993) Ascomycete relationships: dating the
origin of asexual lineages with 18S ribosomal RNA sequence data. In:
Reynolds DR and Taylor JW (eds) The Fungal Holomorph: Mitotic,
Meiotic, and Pleomorphic Speciation in Fungal Systematics, pp. 6778.
Wallingford, UK: CAB International.

10

Lacey J and Crook B (1988) Fungal and actinomycete spores as

pollutants of the workplace and occupational allergens. Annals of
Occupational Hygiene 32: 515533.
Pirozynski KA (1976) Fossil fungi. Annual Review of Phytopathology 14:
237246.

Further Reading
Braunwald E, Isselbacher KJ, Petersdorf RG et al. (eds) (1987)
Harrisons Principles of Internal Medicine. New York: McGraw Hill.
Duddington CL (1973) Zoopagales. In: Ainsworth GC, Sparrow FK and
Sussman AS (eds) The Fungi An Advanced Treatise, vol. 4B, pp. 231
234. New York: Academic Press.
Fraser CM (ed.) (1986) The Merck Veterinary Manual a Handbook of
Diagnosis, Therapy, and Disease Prevention and Control for the
Veterinarian, 6th edn. Rahway, NJ: Merck.
Hawksworth DL, Kirk PM, Sutton BC and Pegler DN (eds) (1995)
Ainsworth and Bisbys Dictionary of the Fungi, 8th edn. Wallingford:
CAB International.
Hesseltine CW (1965) A millennium of fungi, food, and fermentation.
Mycologia 57: 149197.
Hesseltine CW and Ellis JJ (1973) Mucorales. In: Ainsworth GC,
Sparrow FK and Sussman AS (eds) The Fungi An Advanced Treatise,
vol. 4B, pp. 187217. New York: Academic Press.
Lacey J and Crook B (1988) Fungal and actinomycete spores as
pollutants of the workplace and occupational allergens. Annals of
Occupational Hygiene 32: 515533.
Lichtwardt RW (1985) The Trichomycetes: Fungal Associates of
Arthropods. New York: Springer-Verlag.
Moore-Landecker E (1996) Fundamentals of the Fungi, 4th edn. Upper
Saddle River, NJ: Prentice-Hall.
Prescott SC and Dunn CG (1959) Industrial Microbiology, 3rd edn. New
York: Mc Graw-Hill.
Waterhouse GM (1973) Entomophthorales. In: Ainsworth GC, Sparrow
FK and Sussman AS (eds) The Fungi An Advanced Treatise, vol. 4B,
pp. 219229. New York: Academic Press.
Yokotsuka T (1991) Proteinaceous fermented food and condiments
prepares with koji molds. In: Arora DK, Mukerji KG and Marth EH
(eds) Handbook of Applied Mycology, vol. 3: Foods and Feeds, pp. 329
373. New York: Marcel Dekker.