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Introduction to Fungi, Third Edition

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PYTHIALES<br />

97<br />

Fig 5.15 Pythium mycelium in the rotting tissue of a cress<br />

seedling hypocotyl. Note the spherical sporangium initial and<br />

the absence of haus<strong>to</strong>ria.<br />

coenocytic, with typically granular cy<strong>to</strong>plasmic<br />

contents (Fig. 5.15). At first there are no septa,<br />

but later cross walls may cut off empty portions<br />

of hyphae. Thick-walled chlamydospores may<br />

also be formed. There are no haus<strong>to</strong>ria.<br />

Several species are known <strong>to</strong> cause damping<br />

off, e.g. P. debaryanum and, perhaps more<br />

frequently, P. ultimum. Pythium aphanidermatum<br />

is associated with stem rot and damping off of<br />

cucumber, and the fungus may also cause rotting<br />

of mature cucumbers. Pythium mamillatum<br />

causes damping off of mustard and beet seedlings<br />

and is also associated with root rot in Viola.<br />

Many Pythium spp. have a very wide host range;<br />

e.g. P. ultimum parasitizes over 150 plant species<br />

belonging <strong>to</strong> many different families (Middle<strong>to</strong>n,<br />

1943; Hendrix & Campbell, 1973). Far from<br />

parasitizing only plant roots, several soil-borne<br />

species, e.g. P. oligandrum, P. acanthicum and<br />

P. nunn, are capable of attacking hyphae of<br />

filamen<strong>to</strong>us fungi, including plant-pathogenic<br />

species and even other Pythium spp. (Foley &<br />

Deacon, 1986b; Deacon et al., 1990). Attack may<br />

be mediated by the secretion of wall-degrading<br />

b-1,3-glucanase, chitinase and cellulase, or by<br />

inducing the host <strong>to</strong> undergo au<strong>to</strong>lysis (Elad<br />

et al., 1985; Laing & Deacon, 1991; Fang & Tsao,<br />

1995). In contrast <strong>to</strong> plant-pathogenic Pythium<br />

spp., the mycoparasitic species require thiamine<br />

for growth and are unable <strong>to</strong> utilize inorganic<br />

nitrogen sources. These deficiencies may<br />

explain their mycoparasitic habit (Foley &<br />

Deacon, 1986a). Other species of Pythium parasitize<br />

freshwater and marine algae (Kerwin et al.,<br />

1992).<br />

The taxonomy of Pythium is somewhat<br />

confused at present due <strong>to</strong> the existence of<br />

numerous synonyms. Including a few varieties,<br />

Dick (2001a) listed 129 names in current use.<br />

Since the morphological characteristics traditionally<br />

used for diagnosis can be variable, the<br />

delimitation of species and their assignment <strong>to</strong><br />

the genus Pythium will have <strong>to</strong> await the results<br />

of detailed molecular phylogenetic analyses<br />

which are in progress (Matsumo<strong>to</strong> et al., 1999;<br />

Lévesque & de Cock, 2004). Keys and descriptions<br />

have been published by Waterhouse (1967,<br />

1968), van der Plaats-Niterink (1981) and Dick<br />

(1990b).<br />

Asexual reproduction<br />

The mycelium within the host tissue or in<br />

culture usually produces sporangia, but their<br />

form varies. In some species, e.g. P. gracile,

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