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Introduction to Fungi, Third Edition

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232 ASCOMYCOTA (ASCOMYCETES)<br />

Fig 8.5 Two basic modes of development during<br />

conidiogenesis of a hyphal apex (a): blastic (b) and thallic (c).<br />

From de Hoog et al. (2000a), with kind permission of<br />

Centraalbureau voor Schimmelcultures.<br />

state of Discosphaerina; see Fig. 17.25). In certain<br />

yeasts including Saccharomyces, the blastic development<br />

of new daughter cells is known as<br />

budding (see Fig. 10.3). Two kinds of blastic<br />

development have been distinguished:<br />

1. Holoblastic. All the wall layers of the<br />

conidiogenous cell contribute <strong>to</strong> the wall of<br />

the newly formed conidium (see Fig. 8.6b).<br />

Aureobasidium pullulans (Fig. 17.25) and Tricladium<br />

splendens (conidial Hymenoscyphus; see Fig. 25.12)<br />

are examples. In some genera with dark (i.e.<br />

melanized), relatively thick-walled conidiophores<br />

such as Stemphylium and Alternaria (anamorphs<br />

of Pleospora), the conidia develop holoblastically,<br />

but a narrow channel persists in the wall of<br />

the conidiogenous cell through which cy<strong>to</strong>plasm<br />

had passed as the spore expanded. This type of<br />

development has been described as porogenous<br />

(Luttrell, 1963) or tretic (Ellis, 1971a) and the<br />

conidia are sometimes termed porospores or<br />

poroconidia (see Figs. 17.10 17.13; Carroll &<br />

Carroll, 1971; Ellis, 1971b).<br />

2. Enteroblastic. The wall of the conidiogenous<br />

cell is rigid and breaks open. The initial of<br />

the conidium is pushed through the opening<br />

and is surrounded by a newly formed wall<br />

(Fig. 8.6c). Two types of enteroblastic development<br />

have been distinguished, phialidic and<br />

annellidic (see Fig. 8.7). In phialidic development<br />

a basipetal succession of conidia (phialospores,<br />

phialoconidia) develops from a specialized conidiogenous<br />

cell, the phialide (Gr. diminutive of<br />

phialis ¼ flask), usually shaped like a bottle<br />

with a narrow neck. Phialides are formed singly<br />

Fig 8.6 Two alternative types of conidiogenesis starting from<br />

an undifferentiated hyphal apex (a). In holoblastic<br />

conidiogenesis (b), the entire wallbecomes inflated <strong>to</strong> form the<br />

conidium initial. In enteroblastic conidiogenesis (c), the<br />

conidium initial develops through a hole in the rigid outer wall.<br />

From de Hoog et al. (2000a), with kind permission of<br />

Centraalbureau voor Schimmelcultures.<br />

or in clusters at the tip of a conidiophore or,<br />

more rarely, laterally. There may be one or<br />

several nuclei in a phialide. As shown in Fig. 8.8<br />

for Thielaviopsis basicola, the initial of the firstformed<br />

phialoconidium is surrounded by the<br />

apical wall of the phialide and is, in reality,<br />

holoblastic. The phialide wall breaks transversely<br />

near its tip and the first conidium, surrounded<br />

by a newly formed wall and capped by the wall<br />

from the broken tip of the phialide, is pushed<br />

out (Hawes & Beckett, 1977; Ingold, 1981). The<br />

new wall material which encases the phialoconidium<br />

is secreted in the form of a cylinder from<br />

the surface of the cy<strong>to</strong>plasm deep within the<br />

phialide, a process known as ring wall building<br />

(Minter et al., 1983a). Before the conidium is<br />

extruded, a septum develops within the phialide<br />

below its neck, at the base of the conidium.<br />

The upper part of the wall of the now open-ended<br />

phialide persists as a small collar, the collarette.<br />

The nucleus or nuclei within the phialide<br />

continue <strong>to</strong> divide mi<strong>to</strong>tically. A second conidium<br />

develops below the first, and is surrounded<br />

by newly secreted wall material. This conidium is<br />

also cut off by a septum and pushed out. Part of<br />

the newly secreted wall material may persist<br />

around the inside of the neck of the phialide as<br />

periclinal thickening. The process is repeated<br />

so that many phialoconidia may develop from<br />

a single phialide. In phialides which have<br />

developed several conidia, the periclinal

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