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Introduction to Fungi, Third Edition

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UREDINALES: THE RUST FUNGI<br />

611<br />

Fig 22.1 The life cycle of Puccinia graminis, with its heterokaryotic phase on cereals and the homokaryotic stage on barberry<br />

(Berberis).The different ways in which the five spore stages are released and dispersed are indicated.Teliospores overwinter after<br />

nuclear fusion, i.e. as diploid cells. During basidiosporogenesis, meiosis is followed by a mi<strong>to</strong>tic division so that each basidiospore is<br />

a homokaryon containing two nuclei of the same mating type.Open and closed circles represent haploid nuclei of opposite mating<br />

type; diploid nuclei are larger and half-filled. Key events in the life cycle are plasmogamy (P), karyogamy (K) and meiosis (M).<br />

Each cell contains two haploid nuclei of opposite<br />

mating-type which fuse <strong>to</strong> form a diploid nucleus<br />

(see Fig. 22.2c). The teliospore overwinters in the<br />

diploid state, firmly attached <strong>to</strong> the plant tissue<br />

on which it was produced. In spring, meiosis<br />

occurs while each teliospore cell emits a germ<br />

tube which becomes transversely septate and<br />

forms the promycelium or metabasidium. Each<br />

compartment of the metabasidium produces one<br />

basidiospore which initially contains one haploid<br />

nucleus. This commonly divides so that the<br />

mature basidiospore often contains two genetically<br />

identical haploid nuclei and is thus a<br />

dikaryotic homokaryon (Anikster, 1983). Basidiospores<br />

are actively liberated by the surface<br />

tension catapult mechanism involving Buller’s<br />

drop as described before for other basidiomycetes<br />

(Section 18.5).<br />

The basidiospores of P. graminis are unable <strong>to</strong><br />

infect the cereal host but will infect the alternate<br />

host, barberry (Berberis vulgaris). The resulting<br />

primary mycelium is haploid, homo- and monokaryotic.<br />

At the upper (adaxial) surface of the<br />

host leaf, the primary mycelium forms a flaskshaped<br />

fructification known as a spermogonium.<br />

Within the mesophyll, knots of hyphae<br />

form a pro<strong>to</strong>-aecium but do not develop further<br />

at this stage. Minute uninucleate spermatia<br />

are produced from annellide-like structures<br />

(Littlefield & Heath, 1979) within the main body<br />

of the spermogonium, and they aggregate within<br />

a rim of hairs (periphyses) at the spermogonial

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