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Introduction to Fungi, Third Edition

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206 ZYGOMYCOTA<br />

Fig 7.36 Basidiobolusranarum. (a) Stages in the<br />

development of endospores by primary<br />

ballis<strong>to</strong>conidia. Above: cy<strong>to</strong>plasmic contents cleaving<br />

<strong>to</strong> form two pro<strong>to</strong>plasts.Centre: sporangium<br />

containing over a dozen sporangiospores. Below:<br />

sporangium showing breakdown of sporangium wall<br />

(traced from Dykstra,1994). (b) Successive cleavage<br />

of pro<strong>to</strong>plasts from a primary ballis<strong>to</strong>sporic conidium<br />

placed on a rich agar medium <strong>to</strong> form the‘Palmella’<br />

stage. (c) Germination of a primary ballis<strong>to</strong>conidium<br />

<strong>to</strong> form a uninucleate secondary capilliconidium with<br />

a terminal beak which has extruded a sticky hap<strong>to</strong>r.<br />

(d) Capilliconidium which has divided <strong>to</strong> produce<br />

several endospores, some of which have been<br />

released following breakdown of its wall.One of the<br />

endospores is germinating. Scale bars: (a,b) ¼ 20 mm,<br />

(c,d) ¼ 12.5 mm.<br />

Malloch, 1989). Mites are ingested by beetles, the<br />

main vec<strong>to</strong>rs of B. ranarum, but other insects,<br />

spiders, millipedes, woodlice, worms and snails<br />

may also acquire conidia. These are ingested by<br />

vertebrate insectivores. Within the vertebrate<br />

gut, endospores are released from ballis<strong>to</strong>sporic<br />

conidia and capilliconidia.<br />

Endospores germinate in the vertebrate gut<br />

<strong>to</strong> form spherical, large, uninucleate, hyaline<br />

cells up <strong>to</strong> 20 mm. These were called the ‘Darm-<br />

Form’ (gut-stage) of B. ranarum by Levisohn (1927)<br />

who showed that a single ingested primary<br />

conidium can give rise <strong>to</strong> 50 60 division<br />

products (meris<strong>to</strong>spores) forming gut-stage cells<br />

(Fig. 7.34a). The concentration of Basidiobolus<br />

propagules builds up in the guts of the vertebrate<br />

vec<strong>to</strong>rs, and the fungus can be isolated<br />

from faeces of lizards up <strong>to</strong> 18 days after the<br />

animals are deprived of infected prey (Coremans-<br />

Pelseneer, 1973; Okafor et al., 1984). The gut-stage<br />

cells are voided with the faeces and can survive<br />

for several months under dry conditions. Under<br />

moist warm conditions they germinate <strong>to</strong> form a<br />

mycelium from which ballis<strong>to</strong>sporic conidiophores<br />

develop, whereas on certain media they<br />

enlarge and their contents undergo successive<br />

binary fission <strong>to</strong> form globose thick-walled cells.<br />

This state is sometimes termed the ‘Palmella’<br />

state (Fig. 7.36b) because of its superficial<br />

resemblance of a genus of green algae.

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