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Introduction to Fungi, Third Edition

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350 HYMENOASCOMYCETES: PYRENOMYCETES<br />

Fig12.24 Important metabolites from<br />

Clavicipitales. (a) Cyclosporin A, produced by<br />

Tolypocladium inflatum.This is used extensively as<br />

an immunosuppressive drug, e.g. after organ<br />

transplantations. (b) Cephalosporin C,<br />

a b-lactam antibiotic with activity against<br />

Gram-positive and Gram-negative bacteria.<br />

and sits like a cap over it. The ovary, which would<br />

normally develop in<strong>to</strong> a caryopsis filled with<br />

grain, becomes replaced by fungal tissue. For this<br />

reason the disease caused by C. purpurea has been<br />

termed a replacement disease (Luttrell, 1980).<br />

The fungal structure which develops is considerably<br />

longer than the ovary which it replaces and<br />

it differentiates in<strong>to</strong> a sclerotium up <strong>to</strong> 3 cm in<br />

length, the foot of which continues <strong>to</strong> obtain<br />

nutrients via the vascular connection in the host<br />

rachilla. The sclerotium is made up of three<br />

distinct layers: a thin purplish-brown rind, a<br />

discontinuous layer of mealy white tissue, and<br />

a central layer of translucent gelatinous tissue<br />

(Luttrell, 1980).<br />

Ergot is the French name for a cock’s spur,<br />

referring <strong>to</strong> the curved, banana-shaped sclerotia<br />

which project from the inflorescence of infected<br />

grasses and cereals in late summer (Fig. 12.26a).<br />

The sclerotia fall <strong>to</strong> the ground and overwinter<br />

near the surface of the soil. They need a period of<br />

low temperature before they can develop further.<br />

A chilling period of 0°C for at least 25 days<br />

is optimal for further development. The main<br />

reserve substance of the sclerotium is lipid,<br />

which may account for 50% of the dry weight.<br />

It is likely that the chilling period is necessary<br />

before enzymes capable of mobilizing the lipid<br />

reserves develop (Cooke & Mitchell, 1970).<br />

Sclerotia do not remain viable in soil in the<br />

field for more than a few months, often being<br />

invaded by fungi, bacteria, mites and insects<br />

(Cunfer & Seckinger, 1977). The following<br />

summer, sclerotia develop one or more perithecial<br />

stromata (clavae) about 1 2 cm high, shaped<br />

like miniature drumsticks (Fig. 12.26c). The<br />

perithecial stromata are positively pho<strong>to</strong>tropic<br />

(Hadley, 1968).<br />

The enlarged spherical head or capitulum<br />

contains a number of perithecia which are embedded<br />

in the stroma, each surrounded by a<br />

distinct perithecial wall (Fig. 12.27a). The cy<strong>to</strong>logical<br />

details of perithecial development have<br />

been studied in C. purpurea by Killian (1919) and<br />

in C. microcephala (regarded by some as a form of<br />

C. purpurea) by Kulkarni (1963). In the outer layers<br />

of the head of the perithecial stroma, clubshaped<br />

multinucleate antheridia and ascogonia<br />

undergo plasmogamy. Ascogenous hyphae made<br />

up of predominantly binucleate segments<br />

develop from the base of the ascogonium, and<br />

the tips of the ascogenous hyphae form croziers

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