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Introduction to Fungi, Third Edition

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422 HYMENOASCOMYCETES: PEZIZALES (OPERCULATE DISCOMYCETES)<br />

Fig14.6 Ascobolus furfuraceus. (a) Group of asci and<br />

paraphyses.One ascus is mature and contains<br />

purple-pigmented ascospores. (b) The same ascus as<br />

shown in a after discharge.The ascus has decreased in size<br />

during discharge. Note the operculum. (c) Arthrospores<br />

(oidia) developed in five-day-old culture. (d) Coiled<br />

ascogonium formed in a single ascospore culture within<br />

48 h of adding oidia of the opposite mating type.<br />

The trichogyne of the ascogonium has grown <strong>to</strong>wards<br />

the oidium and has fused with it.<br />

14.5.2 Apothecium development<br />

The development of apothecia of A. furfuraceus<br />

has been studied by various authors, including<br />

Wells (1970, 1972) and O’Donnell et al. (1974).<br />

The fertilized ascogonium becomes surrounded<br />

by sheath hyphae which develop from the<br />

ascogonial stalk, and the paraphyses and excipular<br />

tissues develop from the sheath hyphae.<br />

The ascogonium gives rise <strong>to</strong> numerous ascogenous<br />

hyphae. Van Brummelen (1967) has<br />

distinguished two kinds of ascocarp development<br />

in Ascobolus. In gymnohymenial forms,<br />

the hymenium is exposed from the beginning<br />

until the maturation of the asci. In cleis<strong>to</strong>hymenial<br />

forms the hymenium is enclosed during<br />

its early development. Ascobolus furfuraceus and<br />

A. immersus are examples of cleis<strong>to</strong>hymenial<br />

development.<br />

14.5.3 Ascosporogenesis<br />

The details of ascosporogenesis have been<br />

studied in A. immersus by Wu and Kimbrough<br />

(1992) and in A. stic<strong>to</strong>ideus by Wu and Kimbrough<br />

(2001). Although A. immersus has smooth ascospores<br />

and A. stic<strong>to</strong>ideus has ornamented spores,<br />

development is very similar. As in most ascomycetes,<br />

invagination of the ascus vesicle encloses<br />

the young ascospores by a two-layered sporedelimiting<br />

membrane. The inner layer forms the<br />

plasmalemma of the ascospore, whilst the outer<br />

membrane forms a perisporic sac. The primary<br />

spore wall is laid down between these two<br />

membranes. It consists of an electron-transparent<br />

endospore and a laminated epispore. The<br />

perisporic sac is variable in thickness and<br />

projects in<strong>to</strong> the epiplasm of the ascus.<br />

At points of contact with the perisporic sac

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