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Introduction to Fungi, Third Edition

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76 STRAMINIPILA: OOMYCOTA<br />

Fig 5.1 Asexual reproductive stages in Saprolegnia.<br />

(a) Auxiliary (primary) zoospore. (b) Principal<br />

(secondary) zoospore. Schematic drawings, based<br />

partly on Dick (2001a).<br />

rather than lamellate cristae (see Fig. 4.1). The<br />

vacuolar system of Oomycota is also unusual in<br />

containing dense-body vesicles or ‘fingerprint<br />

vacuoles’ (see Fig. 5.24b) which consist of deposits<br />

of a phosphorylated b-(1,3)-glucan polymer, mycolaminarin.<br />

Mycolaminarin may serve as a s<strong>to</strong>rage<br />

compound for carbohydrates as well as phosphate<br />

(Hemmes, 1983), and the polyphosphate<br />

s<strong>to</strong>rage deposits which are typically found within<br />

vacuoles of true <strong>Fungi</strong> are absent from vacuoles<br />

of Oomycota (Chilvers et al., 1985). Apart from<br />

that, however, vacuoles of Oomycota share many<br />

features with those of true <strong>Fungi</strong>, including the<br />

membranous continuities which often link adjacent<br />

vacuoles and provide a means of transport<br />

by peristalsis (Rees et al., 1994; see Fig. 1.9).<br />

Cy<strong>to</strong>plasmic glycogen granules, which are one<br />

of the major carbohydrate s<strong>to</strong>rage sites in<br />

Eumycota, are absent from hyphae of Oomycota<br />

(Bartnicki-Garcia & Wang, 1983).<br />

5.1.2 The zoospore<br />

The Oomycota are characterized by motile asexual<br />

spores (zoospores) which are produced in<br />

spherical or elongated zoosporangia. They are<br />

heterokont, possessing one straminipilous and<br />

one whiplash-type flagellum. Two types of zoospore<br />

may be produced and, if so, the auxiliary<br />

zoospore is the first formed. It is grapeseedshaped,<br />

with both flagella inserted apically<br />

(Fig. 5.1a), and it encysts soon after its formation.<br />

Encystment is by withdrawal of the flagella,<br />

so that a tuft of tripartite tubular hairs (TTHs;<br />

see p. 68) is left behind on the surface of the<br />

developing cyst (Dick, 2001b). The cyst germinates<br />

<strong>to</strong> give rise <strong>to</strong> the principal zoospore, which is<br />

by far the more common type and also the more<br />

vigorous swimmer. This typical and readily recognized<br />

oomycete zoospore is uniform in appearance<br />

across the phylum (Lange & Olson, 1983;<br />

Dick, 2001a). In species lacking auxiliary zoospores,<br />

the principal zoospore is usually produced<br />

directly from a sporangium. It is kidney-shaped,<br />

with the flagella inserted laterally in a kine<strong>to</strong>some<br />

boss which in turn is located within the<br />

lateral groove (Fig. 5.1b). Encystment is initiated<br />

by the shedding, rather than withdrawal, of the<br />

flagella; no tufts of TTHs are left on the cyst<br />

surface (Dick, 2001a). Fascinating insights in<strong>to</strong><br />

the cy<strong>to</strong>logy of zoospore encystment have been

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