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Introduction to Fungi, Third Edition

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240 ASCOMYCOTA (ASCOMYCETES)<br />

Fig 8.12 Types of ascus dehiscence. (a) Pro<strong>to</strong>tunicate ascus; the wall dissolves <strong>to</strong> release the ascospores passively. (b,c) Operculate<br />

asci before and after discharge; the ascus opens by means of a lid or operculum. (d) Discharged inoperculate ascus which has opened<br />

through a pore. (e,f) Rostrate ascus as seen in Physcia. In (e) a thickened part of the upper wall of the ascus is being extruded and<br />

is visible in a discharged ascus (f) as an extension of the inner part of the ascus wall, the rostrum. (g) Discharged bilabiate ascus<br />

showing the longitudinal slit by which the ascus opens. (h,i) Bitunicate ascus before and after the first stage of spore release.<br />

Rupture of the ec<strong>to</strong>tunica has allowed the endotunica <strong>to</strong> expand. (e,f) after Honegger (1978).<br />

ascus mother cell. During this stage the outer<br />

layers of the wall making up the ec<strong>to</strong>tunica<br />

(¼ ec<strong>to</strong>ascus) are deposited. In the second stage,<br />

secondary wall layers making up the endotunica<br />

(¼ endoascus) are laid down within the primary<br />

wall. The development of bitunicate asci has<br />

been studied by Reynolds (1971) and by Parguey-<br />

Leduc and Janex-Favre (1982). At the beginning<br />

of development, asci are surrounded by a single<br />

homogeneous layer which is sometimes granular,<br />

bearing externally a loose network (a fuzzy<br />

coat) of interascal material. The ascus wall<br />

becomes divided in<strong>to</strong> a densely granular external<br />

layer (the ec<strong>to</strong>ascus) and a clearer, but equally<br />

granular, inner layer (the endoascus). A clear<br />

space then separates these two layers. The<br />

granular material of the endoascus rearranges<br />

itself in<strong>to</strong> lines of fibrils at first following a wavy<br />

pattern as seen in transverse sections of developing<br />

asci. Later the fibrils become strongly<br />

folded in<strong>to</strong> pointed zigzag shapes, a development<br />

which progresses from the inside <strong>to</strong>wards the<br />

outside of the endoascus. The folds of the zigzags<br />

are closely pressed against the pointed teeth<br />

which mark out the plasmalemma. The density<br />

of the fibrils increases considerably throughout<br />

the thickness of the endoascus. Finally the two<br />

layers of the ascal wall, separated from each<br />

other by a clear space, appear as a double-layered<br />

ec<strong>to</strong>ascus and a single-layered endoascus within<br />

which the fibrils are strongly pleated in<strong>to</strong><br />

accordion-like folds. The pointed crests of the<br />

pleats lie parallel <strong>to</strong> each other and perpendicular<br />

<strong>to</strong> the plasmalemma of the ascus. The<br />

folding of the layers of the endoascus and<br />

plasmalemma permit the rapid expansion of<br />

the ascus prior <strong>to</strong> spore discharge, i.e. by<br />

providing material which can unfold rapidly.<br />

Towards the tip of the non-discharged ascus<br />

the crests of the pleated folds of the endoascus

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