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Introduction to Fungi, Third Edition

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98 STRAMINIPILA: OOMYCOTA<br />

Fig 5.16 Sporangia and zoospores of Pythium.(a)Pythium debaryanum. Spherical sporangium with short tube and a vesicle<br />

containing zoospores. (b k) Pythiumaphanidermatum. (b) Lobed sporangium showing a long tube and the vesicle, which is beginning<br />

<strong>to</strong> expand. (c g) Further stages in the enlargement of the vesicle, and differentiation of zoospores.Note the transfer of pro<strong>to</strong>plasm<br />

from the sporangium <strong>to</strong> the vesicle in (c).The stages illustrated in (b g) <strong>to</strong>ok place in 25 min. (h) Enlarged vesicle showing the<br />

zoospores. Flagella are also visible. (i) Zoospores. (j) Encystment of zoospore showing a shed flagellum. (k) Germination of<br />

azoosporecyst.(b g) <strong>to</strong> same scale; (a) and (h k) <strong>to</strong> same scale.<br />

the sporangia are filamen<strong>to</strong>us and are scarcely<br />

distinguishable from vegetative hyphae.<br />

In P. aphanidermatum, the sporangia are formed<br />

from inflated lobed hyphae (Fig. 5.16b). In many<br />

species, however, e.g. P. debaryanum, the sporangia<br />

are globose (Fig. 5.16a). A terminal or<br />

intercalary portion of a hypha enlarges and<br />

assumes a spherical shape, then becomes cut<br />

off from the mycelium by a cross wall. The<br />

sporangia contain numerous nuclei. Cleavage of<br />

the cy<strong>to</strong>plasm <strong>to</strong> form zoospores begins in the<br />

sporangium, but is completed within a thinwalled<br />

vesicle which is extruded from the<br />

sporangium. This is a homohylic vesicle because<br />

its glucan wall is continuous with one layer of<br />

the sporangial wall (Dick, 2001a). Within the<br />

sporangium, cleavage vesicles begin <strong>to</strong> coalesce<br />

<strong>to</strong> separate the cy<strong>to</strong>plasm in<strong>to</strong> uninucleate<br />

portions; membrane-bound packets of TTHs<br />

are already present within the cy<strong>to</strong>plasm of the<br />

sporangium. In P. middle<strong>to</strong>nii (Fig. 5.17), the<br />

fascinating process of differentiation from amorphous<br />

cy<strong>to</strong>plasm <strong>to</strong> motile zoospores takes about<br />

30 45 min (Webster, 2006a) and is readily<br />

demonstrated in the labora<strong>to</strong>ry (Weber et al.,<br />

1999). The sporangium is extended in<strong>to</strong> an apical<br />

papilla capped by a mass of fibrillar material<br />

which is lamellate in ultrastructure (Lunney &<br />

Bland, 1976). Shortly before sporangial discharge,<br />

there is an accumulation of cleavage vesicles<br />

behind the apical cap and at the periphery of<br />

the cy<strong>to</strong>plasm close <strong>to</strong> the sporangium wall. The<br />

cleavage vesicles around the sporangial cy<strong>to</strong>plasm<br />

discharge their contents <strong>to</strong> form a loose,<br />

fibrous interface between the cy<strong>to</strong>plasm and the<br />

sporangial wall.<br />

Discharge of the sporangium occurs by the<br />

formation of a thin-walled vesicle at the tip of<br />

the papilla from the fibrillar material of the<br />

apical cap, and the partially differentiated<br />

zoospore mass is extruded in<strong>to</strong> it. The movement<br />

of the cy<strong>to</strong>plasm from the sporangium in<strong>to</strong> the

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