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Introduction to Fungi, Third Edition

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CERATOBASIDIALES<br />

595<br />

affected (Sneh et al., 1991), the question of the<br />

boundaries of biological species remains unanswerable<br />

at present.<br />

The basidiocarps of Cera<strong>to</strong>basidiales are thin,<br />

gelatinous and often resupinate. Their formation<br />

can sometimes be induced by covering an agar<br />

culture with soil (Warcup & Talbot, 1966). Hyphal<br />

tips due <strong>to</strong> develop in<strong>to</strong> a probasidium are<br />

generally binucleate, and karyogamy is followed<br />

swiftly by meiosis. During the later stages of<br />

meiosis, four prominent sterigmata referred <strong>to</strong><br />

as epibasidia are formed (Wells & Bandoni, 2001),<br />

but septa are not laid down, even in the mature<br />

basidium (Fig. 21.1a). The basidiospores are uninucleate<br />

and frequently germinate by repetition<br />

<strong>to</strong> produce uninucleate ballis<strong>to</strong>conidia. Eventually,<br />

hyphal germination occurs. Mating systems<br />

in the Cera<strong>to</strong>basidiales are not well unders<strong>to</strong>od,<br />

and several species may be homothallic.<br />

The basidium of the Tulasnellales differs in<br />

that the four epibasidia become separated from<br />

the metabasidium by septation after meiosis.<br />

Maturation of the four epibasidia may occur at<br />

different times (Fig. 21.1b).<br />

21.2.1 Rhizoc<strong>to</strong>nia in agriculture<br />

The most important taxon is R. (Moniliopsis) solani<br />

(teleomorph Thanatephorus cucumeris), which causes<br />

a wide array of soil-borne necrotrophic diseases<br />

especially of herbaceous plants including all<br />

kinds of vegetables, rice, turfgrasses, and less<br />

frequently also woody tree hosts (Adam, 1988;<br />

Agrios, 2005). The most common disease is<br />

damping off of seedlings in which the infected<br />

hypocotyl region becomes water-soaked and no<br />

longer provides structural integrity (see p. 95),<br />

leading <strong>to</strong> pre- or post-emergence death of<br />

the seedling. In older plants, infection may not<br />

immediately cover the entire circumference of<br />

the plant stem, so that cankers and girdling of the<br />

stem may result. Roots are also frequently<br />

infected, as are aerial plant organs in contact<br />

with the soil or exposed <strong>to</strong> watersplash from the<br />

soil surface. The fungus can survive in the soil<br />

for several years as small sclerotia about 1 3mm<br />

in diameter. These are often associated with<br />

the debris of host plants killed by the fungus.<br />

Sclerotia are occasionally seen as black scurf on<br />

the surface of pota<strong>to</strong> tubers because they are not<br />

easily removed, even by assiduous washing.<br />

Rhizoc<strong>to</strong>nia diseases are typically most severe<br />

at cool temperatures, presumably because the<br />

seedling stage of host plants is prolonged due<br />

<strong>to</strong> their slow growth. The infection process<br />

is similar <strong>to</strong> that of Gaeumannomyces graminis,<br />

i.e. hyphae branch and build up a cushion-like<br />

compound appressorium on the host plant<br />

surface, from which penetration is achieved.<br />

Fig 21.1 (a) Basidium of Thanatephorus cucumeris<br />

(Cera<strong>to</strong>basidiales). Note the four prominent<br />

epibasidia and the lack of septation within the<br />

basidium. (b) Basidium of Gloeotulasnella cystidiophora<br />

(Tulasnellales) showing various stages of maturity.<br />

The epibasidia are separated from the metabasidium<br />

by septation.Three epibasidia have already<br />

discharged their basidiospore, with one about <strong>to</strong><br />

produce it. (a) redrawn from Warcup and Talbot<br />

(1962) with permission from Elsevier, (b) redrawn<br />

from Wells and Bandoni (2001) with kind permission<br />

of Springer Science and Business Media.

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