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Introduction to Fungi, Third Edition

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282 HEMIASCOMYCETES<br />

form septa and quickly fragment in<strong>to</strong> arthrospores<br />

(Fig. 10.10), giving the colonies a creamy<br />

appearance. The septa have micropores, like<br />

those of true hyphae of C. albicans. Six species<br />

are now known; by far the most important is<br />

G. candidus (anamorph Geotrichum candidum),<br />

formerly known as Galac<strong>to</strong>myces geotrichum (de<br />

Hoog & Smith, 2004). It is a ubiqui<strong>to</strong>us mould<br />

which is common in soil, dairy products, sewage<br />

and other substrata. It is also thought <strong>to</strong> be a<br />

common constituent of the skin and gut flora<br />

of humans and animals, although reports of<br />

it being a human pathogen have generally<br />

remained unsubstantiated (Kwon-Chung &<br />

Bennett, 1992). In addition, G. candidus is a wellknown<br />

cause of post-harvest rot in ripe fruits<br />

and vegetables especially when these are kept<br />

in plastic bags. Infected plant organs become<br />

soft and eventually, upon puncturing, exude<br />

a creamy mass of decaying tissue which has<br />

a sour smell; hence the name ‘sour rot’ (Agrios,<br />

2005).<br />

The fungus grows readily in culture, forming<br />

broad hyphae with finer lateral branches. The<br />

vegetative cells contain 1 4 nuclei. Branching<br />

is of two kinds, pseudodicho<strong>to</strong>mous near the<br />

apex, and lateral immediately behind a septum.<br />

It is from such lateral branches that conidia<br />

develop (Figs. 10.10a,b). Conidiophores are difficult<br />

<strong>to</strong> differentiate from vegetative hyphae.<br />

Prior <strong>to</strong> conidium formation, apical growth of<br />

a hypha ceases, then septa are laid down in the<br />

tip region. The septa are two-ply, and separation<br />

of the two layers making up the septum<br />

leads <strong>to</strong> the disarticulation of the terminal part<br />

of a hypha in<strong>to</strong> cylindrical segments termed<br />

arthrospores or arthroconidia (Cole & Kendrick,<br />

1969b). Conidia of other Galac<strong>to</strong>myces spp. are<br />

virtually indistinguishable from those of<br />

G. candidus.<br />

Galac<strong>to</strong>myces candidus may be homo- or heterothallic,<br />

but the sexual state is not frequently<br />

seen. After mating, fertile hyphae are produced<br />

and gametangia arise in pairs on either side of<br />

a septum, in the broad main hyphae or short<br />

side branches (Figs. 10.10c e). Fusion of the<br />

gametangia gives rise <strong>to</strong> a globose fusion cell<br />

which becomes transformed directly in<strong>to</strong> an<br />

ascus. The ascus contains only a single ascospore<br />

which has two wall layers, a smooth inner layer<br />

and a furrowed outer layer. Each ascospore<br />

contains 1 2 nuclei. Whether and when meiosis<br />

occurs is not yet known.<br />

The Geotrichum arthroconidial state is found<br />

also in the only other genus of the Dipodascaceae,<br />

Dipodascus, which produces multispored<br />

asci with 4 128 spores. A superficially similar<br />

state, Saprochaete, is formed by a genus of phylogenetically<br />

unrelated fungi now called Magnusiomyces.<br />

Species descriptions and a key of<br />

Galac<strong>to</strong>myces, Dipodascus and Magnusiomyces have<br />

been provided by de Hoog and Smith (2004).<br />

10.6 Saccharomycopsis<br />

(Saccharomycopsidaceae)<br />

Saccharomycopsis (formerly Endomycopsis) is a mycelial<br />

yeast which reproduces by buds (blas<strong>to</strong>spores<br />

or yeast cells) and also forms asci parthenogenetically<br />

or following isogamous fusion. About<br />

10 species are known (Kurtzman & Smith, 1998;<br />

Barnett et al., 2000). Saccharomycopsis fibuligera<br />

grows in flour, bread, macaroni and other<br />

starchy substrates, and produces a complex of<br />

numerous active extracellular amylases, an<br />

unusual property in yeasts (Hostinová, 2002).<br />

This has been used <strong>to</strong> develop S. fibuligera as<br />

a food yeast for cattle feed which can be grown<br />

on pota<strong>to</strong> starch processing wastes (Jarl, 1969).<br />

This species is also used extensively for starch<br />

hydrolysis by starter cultures in Far Eastern<br />

fermented food (Beuchat, 1995).<br />

In culture, S. fibuligera may form budding<br />

yeast cells and branched septate hyphae which<br />

produce blas<strong>to</strong>spores laterally and terminally<br />

(Fig. 10.11). Arthrospore formation has also<br />

been demonstrated. Ascus formation in this<br />

homothallic species can be induced by growing<br />

the yeast for a few days on malt extract agar<br />

and transferring it <strong>to</strong> distilled water. The asci<br />

are mostly four-spored, and the spores are<br />

hat-shaped (Fig. 10.11d), having a flange-like<br />

extension of the wall.

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