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Introduction to Fungi, Third Edition

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ASCOBOLUS (ASCOBOLACEAE)<br />

421<br />

Fig14.5 Ascobolus immersus.(a)Apothecium<br />

showing two projecting asci. Immature asci can<br />

be seen below the general level of the surface.<br />

A single projectile consisting of eight adhering<br />

ascospores is shown above the apothecium.<br />

Note the operculum which has also been<br />

projected. (b) Tip of ripe ascus showing the<br />

operculum. (c) Tip of discharged ascus. In this<br />

case the operculum has remained attached <strong>to</strong><br />

the ascus tip.<br />

and a, and fertilization can only occur between<br />

an A ascogonium and an a antheridium, or<br />

vice versa. There is thus a gene for mating<br />

type represented in two idiomorphs A and a,<br />

and incompatibility is controlled by this gene<br />

irrespective of the presence of both types of<br />

sex organ on each strain. There is no morphological<br />

difference between the two different<br />

mating type strains.<br />

A similar situation occurs in A. furfuraceus,<br />

but here there are no antheridia. Instead, each<br />

strain at first produces chains of arthrospores<br />

or oidia (see Fig. 14.6c). The oidia can germinate<br />

<strong>to</strong> form a fresh mycelium, i.e. they can function<br />

asexually as conidia, but they also play a part<br />

in sexual reproduction. Mites and flies may<br />

transport oidia of one strain <strong>to</strong> the mycelium<br />

of the opposite strain, and following this,<br />

apothecia develop. The process of fertilization<br />

has been studied by Bistis (1956, 1957) and Bistis<br />

and Raper (1963). If an A oidium is transferred<br />

<strong>to</strong> an a mycelium, the oidium fails <strong>to</strong> germinate<br />

and within 10 h an ascogonial primordium<br />

appears on the a mycelium (Fig. 14.6d). The<br />

ascogonium consists of a broad coiled base and<br />

a narrow apical trichogyne which shows chemotropic<br />

growth <strong>to</strong>wards the oidium and eventually<br />

fuses with it. There is evidence that this sequence<br />

of events is under hormonal control, and it<br />

has been suggested that a fresh A oidium is not<br />

immediately capable of inducing development<br />

of ascogonial primordia, but must itself at first<br />

be sexually activated by a messenger secreted<br />

by the a mycelium. Following activation, the<br />

oidium can induce ascogonial development.<br />

By substitution experiments, it has been shown<br />

that an A ascogonium can be induced <strong>to</strong> fuse<br />

with an A oidium, i.e. an oidium of the same<br />

mating type, but apothecia fail <strong>to</strong> develop<br />

from such fusions. In compatible crosses, fertile<br />

apothecia develop within about 10 days of fertilization,<br />

each ascus producing four A and four<br />

a spores.<br />

In A. immersus there are no morphologically<br />

distinguishable antheridia and there are<br />

no oidia. When A and a mating type mycelia<br />

are grown <strong>to</strong>gether in culture, multinucleate<br />

ascogonia develop which are fertilized by fusion<br />

with slender multinucleate hyphae of the<br />

opposite strain. Experimentally, fertilization<br />

can also be achieved using homogenized mycelial<br />

fragments of one strain <strong>to</strong> spermatize a<br />

strain of opposite mating type (Lewis & Decaris,<br />

1974).

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