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Introduction to Fungi, Third Edition

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CONIDIUM PRODUCTION IN ASCOMYCETES<br />

231<br />

where the hyphae making up the synnema are<br />

intricately interwoven (see Kirk et al., 2001).<br />

In many ascomycetes the conidiophores develop<br />

on or in a stroma (Gr. stroma ¼ bed, cushion),<br />

an aggregation of pseudoparenchyma<strong>to</strong>us cells.<br />

A good example of a conidial stroma is seen in<br />

the wood-rotting candle-snuff fungus, Xylaria<br />

hypoxylon (see Fig. 12.11a). Here, powdery white<br />

conidia develop at the tips of the branches of<br />

the conidial stroma and, later, asci develop in<br />

flask-shaped perithecia at the base of the old<br />

stroma. The term sporodochium (Gr. spora ¼ a<br />

seed; doche ¼ a receptacle) is used for the<br />

cushion-like conidiomata bearing a layer of<br />

short conidiophores. An example is the conidial<br />

(Tubercularia) state of Nectria cinnabarina (see<br />

Fig. 12.20c).<br />

Another type of conidioma is the acervulus<br />

(Lat. acervulus ¼ a little heap), a saucer-shaped<br />

fructification which may develop inside the<br />

tissues of a host plant or may be superficial.<br />

Subepidermal acervuli develop from a pseudoparenchyma<strong>to</strong>us<br />

stroma, and as the acervulus<br />

matures the overlying epidermis of the host<br />

becomes ruptured <strong>to</strong> expose conidia formed<br />

from conidiogenous cells lining the base of the<br />

saucer. The conidia are held <strong>to</strong>gether in slime<br />

and are chiefly dispersed by rain splash. A good<br />

example of an acervular fungus is Colle<strong>to</strong>trichum<br />

(see Fig. 12.51). The teleomorphs of Colle<strong>to</strong>trichum,<br />

where known, are species of Glomerella, many of<br />

which are serious plant pathogens. In many<br />

ascomycetes and their allies, the conidia are<br />

borne inside flask-shaped conidiomata termed<br />

pycnidia (Gr. diminutive of pyknos ¼ dense,<br />

packed, concentrated). Traditionally, fungi with<br />

pycnidial and acervular states have been grouped<br />

<strong>to</strong>gether in the artificial taxon coelomycetes<br />

(Sut<strong>to</strong>n, 1980), in contrast <strong>to</strong> hyphomycetes in<br />

which the conidiogenous cells are exposed on<br />

single conidiophores or in synnemata, coremia<br />

or sporodochia (see above). Pycnidia may be<br />

superficial or embedded in host tissue. The opening<br />

of the pycnidium is generally by means of<br />

a circular ostiole. Conidia formed from conidiogenous<br />

cells lining the inner wall of the<br />

pycnidium are held <strong>to</strong>gether in slimy masses<br />

which ooze out through the ostiole, sometimes<br />

as spore tendrils. They are generally dispersed<br />

by splash or in water films. In some cases the<br />

pycnidia, instead of producing conidia with an<br />

asexual function, produce spermatia which are<br />

involved in fertilization. Examples of fungi with<br />

pycnidial anamorphs are Lep<strong>to</strong>sphaeria acuta with<br />

a Phoma anamorph (Fig. 17.3), and Phaeosphaeria<br />

nodorum (anamorph Stagonospora nodorum; see<br />

Fig. 17.4).<br />

8.5 Conidium production<br />

in ascomycetes<br />

There are several steps in the production and<br />

release of conidia, namely (1) conidiogenesis,<br />

i.e. conidial initiation; (2) maturation; (3) delimitation;<br />

(4) secession, i.e. separation from<br />

the conidiogenous cell; (5) proliferation of the<br />

conidiogenous cell or conidiophore <strong>to</strong> form<br />

further conidia. Many of the current ideas on<br />

conidiogenesis stem from a seminal paper by<br />

Hughes (1953) based on light microscopy<br />

studies of conidial development in a range of<br />

hyphomycetes. Hughes classified the development<br />

of conidia in a limited number of ways.<br />

His ideas were extended by other workers, and<br />

advances were also made possible by the use<br />

of electron microscopy and time-lapse cinepho<strong>to</strong>micrography<br />

(Cole & Samson, 1979).<br />

An excellent review of these aspects of<br />

conidiogenesis has been written by Cole (1986).<br />

The descriptions which follow are based on the<br />

account by de Hoog et al. (2000a). Conidiogenesis<br />

occurs in two ways which appear <strong>to</strong> be distinct<br />

at first glance: blastic and thallic (see Fig. 8.5).<br />

In reality, when surveying conidium formation<br />

and release in a range of fungi, there is a<br />

continuum of development of which these two<br />

concepts represent extremes (Minter et al., 1982,<br />

1983a,b; Minter, 1984).<br />

8.5.1 Blastic conidiogenesis<br />

The conidium develops by the blowing-out of<br />

the wall of a cell, usually from the tip of a hypha,<br />

sometimes laterally as in Aureobasidium (conidial

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