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Introduction to Fungi, Third Edition

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110 STRAMINIPILA: OOMYCOTA<br />

Fig 5.26 Phy<strong>to</strong>phthora cac<strong>to</strong>rum.<br />

Development of oogonium, antheridium<br />

and oospore. (a) Initials of oogonium<br />

and antheridium. (b) Oogonium and<br />

antheridium grown <strong>to</strong> full size: the<br />

oogonium has about 24 nuclei and the<br />

antheridium about 9. (c) Development<br />

of a septum at the base of each, and<br />

degeneration of some nuclei in each until<br />

the oogonium has 8 or 9 nuclei and the<br />

antheridium 4 or 5. (d) A simultaneous<br />

division of the surviving nuclei in<br />

oogonium and antheridium.The<br />

pro<strong>to</strong>plast has large vacuoles.<br />

(e) Separation of oosphere from<br />

periplasm. Nuclei divide in the periplasm<br />

prior <strong>to</strong> degeneration.The oogonium<br />

presses in<strong>to</strong> the antheridium. (f) Entry of<br />

one antheridial nucleus by a fertilization<br />

tube.The pro<strong>to</strong>plasm and remaining<br />

nuclei of the antheridium degenerate.<br />

(g) Development of oospore wall.<br />

(h) The oospore enters its dormant<br />

period with exospore formed from dead<br />

periplasm, endospore deposited inside<br />

it, and paired nuclei in association but<br />

not yet fused. (a h) are composite<br />

drawings of eight stages in sequence<br />

(after Blackwell,1943).<br />

a resting state. Oospore differentiation proceeds<br />

from the outside inwards (centripetal development).<br />

The oospore has a thin outer wall<br />

(epispore) which is derived from the periplasm<br />

and appears <strong>to</strong> consist of pectic substances. The<br />

inner oospore wall (endospore) is rich in b-1,3-<br />

glucans which form a major s<strong>to</strong>rage reserve and<br />

are mobilized by glucanases just prior <strong>to</strong> germination<br />

(Erwin & Ribeiro, 1996). Within the<br />

developing oospore, the numerous small lipid<br />

droplets coalesce in<strong>to</strong> a few large ones. Lipids are<br />

undoubtedly the major endogenous s<strong>to</strong>rage<br />

reserve in the spores of Oomycota (Dick, 1995)<br />

and many other fungi. Later, the dense body<br />

vesicles which are rich in mycolaminarin and<br />

phosphate fuse <strong>to</strong>gether, giving one large structure,<br />

the ooplast. Like the endospore, the ooplast<br />

is consumed during germination whereas some<br />

lipid droplets are saved and are translocated in<strong>to</strong><br />

the germ tube (Hemmes, 1983). Considering their<br />

thick walls and abundant s<strong>to</strong>rage reserves, it is<br />

not surprising that oospores are the longest-lived

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