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Introduction to Fungi, Third Edition

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HYPOCREALES<br />

345<br />

Fig12.21 Nectria mammoidea. (a) Ascus;<br />

note the apical apparatus.<br />

(b) Cylindrocarpon-type conidiophore with<br />

phialides and conidia.<br />

from such pustules (see Jenkinson & Parry,<br />

1994). In culture, greasy accumulations of macroconidia<br />

are termed pionnotes. Survival in soil is<br />

probably in the form of dormant chlamydospores<br />

which may be formed as a result of energy<br />

deprivation or in response <strong>to</strong> bacterial secretions.<br />

They are thick-walled and develop<br />

by modification of segments of the vegetative<br />

mycelium or by enlargement and modification<br />

of a segment of a macroconidium (Schippers &<br />

van Eck, 1981) (Figs. 12.22c,d). Chlamydospores<br />

are induced <strong>to</strong> germinate in the presence of root<br />

secretions (for references see Griffiths, 1974).<br />

One of the most commonly isolated species<br />

from soil is F. oxysporum, which often grows in<br />

association with roots. Many isolates are nonpathogenic<br />

but this fungus may also be a serious<br />

plant pathogen, reported from a very wide range<br />

of plant hosts. Although no sexual state is<br />

known, molecular studies show that it is monophyletic<br />

with the Gibberella Fusarium complex<br />

and can be regarded as an asexual Gibberella<br />

(Samuels et al., 2001). In this species over 80<br />

formae speciales characteristic of different host<br />

genera have been recognized, in addition <strong>to</strong> a<br />

large number of vegetative compatibility types<br />

and races (Gordon, 1993; Gordon & Martyn,<br />

1997). The large amount of genetic variation in<br />

F. oxysporum in the absence of conventional<br />

sexual reproduction is possibly explained by<br />

the demonstration of parasexual recombination<br />

even between members of different vegetative<br />

compatibility groups (Molnár et al., 1990).<br />

Fusarium wilts<br />

As can be seen from Table 12.3, formae speciales of<br />

F. oxysporum cause wilt diseases of numerous crop<br />

plants. The range of affected hosts is so wide that<br />

it is easier <strong>to</strong> mention those plants unaffected by<br />

Fusarium wilts, i.e. grasses and many tree species.<br />

Although F. oxysporum has no known sexual state,<br />

strains of this species complex show huge<br />

genetic variation which may come about by the<br />

presence of transposable elements (Daboussi &<br />

Capy, 2003) and the parasexual cycle (Teunissen<br />

et al., 2002). The variability of F. oxysporum is<br />

such that a given disease, e.g. Panama wilt of<br />

banana, is caused by several unrelated strains<br />

(K. O’Donnell et al., 1998), and that strains with<br />

different host specificities may be closely related<br />

(Hua-Van et al., 2001).<br />

Wilting is associated with the presence of<br />

fungal hyphae in the xylem vessels. Fusarium<br />

oxysporum can persist in the soil as chlamydospore<br />

inoculum for many years and infects the<br />

roots of hosts by direct penetration and

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