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membrane (Fig. 12.10). The parasite draws nutrients
from the host hyphae, and uses these nutrients to
produce sporulating structures on the host colony.
Often this type of parasitism causes little damage, as
long as the host fungus has an adequate food supply.
Biotrophic mycoparasitism similar to that in Fig. 12.10
is found in several Zygomycota (Piptocephalis,Dispira,
and Dimargalisspp.) that have elongated, few-spored
sporangia (merosporangia; see Fig. 2.11). With only few
exceptions, these fungi parasitize other Zygomycota such
asMucorandPilairaon dung or in soil. Most of these
biotrophic mycoparasites can be grown in laboratory
media containing extracts of host or nonhost hyphae.
The need for hyphal extracts can be replaced by
relatively high concentrations of vitamins (especially
thiamine) and amino acids, and by providing glycerol
instead of glucose as the carbon source. These nutrients
could be expected to occur in the host hyphae.
The haustorial biotrophs seem to depend entirely on
their fungal hosts in nature. Their spores are triggered
to germinate near host hyphae, and the germ-tubes

show pronounced tropism towards the host. Then the
germ-tube tip produces an appressorium on the host
surface and a penetration peg enters the host to form
a haustorium. The mycoparasite Piptocephalis virginiana
shows evidence of specific recognition in the infection
process (Manocha & Chen 1990). It infects only some
members of the order Mucorales, and the walls of
these hosts were found to have two surface-located
glycoproteins. Removal of these glycoproteins by
treating hyphae with NaOH or proteinase led to
impaired attachment and appressorium formation. The
treated hyphae (with their glycoproteins removed)
were found to bind lectins that recognize fucose,
N-acetylgalactosamine, and galactose, and this binding
pattern of the treated hyphae was identical to that of
nonhost hyphae. However the untreatedhost hyphae
did not bind these lectins, so it is suggested that the
glycoproteins that occur normally on host hyphae
cover the fucose,N-acetylgalactosamine, and galactose
residues (which somehow interfere with infection), and
enable the parasite to attach and infect. In support

FUNGAL INTERACTIONS 245

Fig. 12.10Electron micrograph of an
appressorium (ap) and a branched
haustorium of the mycoparasite
Piptocephalis unispora(Zygomycota) in
a fungal host, Cokeromyces recurvatus.
hw =host wall; ol and il =outer layer
and inner layer of the Piptocephalis
wall. The haustorium is surrounded
by a continuous membrane (the extra-
haustorial membrane, labelled e).
(Courtesy of P. Jeffries; from Jeffries &
Young 1976.)

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