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Introduction to Fungi, Third Edition

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OTHER CEREAL RUSTS<br />

627<br />

race characterization has been written by Roelfs<br />

(1984).<br />

The serious race 15B epidemics of the early<br />

1950s (see Fig. 22.12) resulted in the replacement<br />

of the then prevalent cultivars by others such as<br />

‘Selkirk’, which carried several resistance genes<br />

and remained resistant <strong>to</strong> P. graminis f. sp. tritici<br />

in North America for several decades (Roelfs,<br />

1985). Several wheat varieties more recently<br />

released have also retained their resistance for<br />

many years in the field (Roelfs, 1984; Dyck &<br />

Kerber, 1985), so that no major stem rust<br />

epidemic has occurred in the USA during the<br />

past 50 years. This success is probably due <strong>to</strong> two<br />

main reasons, namely the combination of several<br />

resistance genes in one cultivar which the rust<br />

fungus is apparently unable <strong>to</strong> overcome, and<br />

the stabilization of rust races by the elimination<br />

of sexual reproduction as a consequence of the<br />

barberry eradication campaign.<br />

22.4 Other cereal rusts<br />

Although potentially the most damaging cereal<br />

rust, P. graminis has lost much of its menace<br />

for reasons discussed in the preceding section.<br />

Several other rust species now cause more<br />

serious crop losses than P. graminis. These are<br />

briefly discussed below. Cereal rusts are primarily<br />

controlled by breeding of resistant cereal<br />

cultivars, and this is an ongoing battle against<br />

new races which, once arisen, are capable of<br />

spreading rapidly and on a global scale (Chen,<br />

2005; Kolmer, 2005). Chemical control by the<br />

application of fungicides is practised <strong>to</strong> protect<br />

crops grown for seed production, and <strong>to</strong> control<br />

severe outbreaks of rust if resistance fails. <strong>Fungi</strong>cides<br />

used include the systemic strobilurin-type<br />

compounds and the ergosterol biosynthesisinhibiting<br />

triazoles (see Fig. 13.15), as well as<br />

various protectant compounds.<br />

22.4.1 Puccinia triticina (brown leaf<br />

rust of wheat)<br />

Puccinia triticina was formerly named P. recondita<br />

f. sp. tritici, but Zambino and Szabo (1993) and<br />

Anikster et al. (1997) have shown that it is not<br />

closely related <strong>to</strong> P. recondita, which infects rye<br />

but is only of minor significance. Puccinia triticina<br />

has displaced P. graminis as the economically<br />

most important rust of wheat especially in North<br />

America and Eastern Europe (Samborski, 1985).<br />

This species is macrocyclic, with Thalictrum speciosissimum<br />

(¼ T. flavum; Ranunculaceae) serving as<br />

the alternate host, although it seems <strong>to</strong> survive<br />

mainly in the uredinial state. Its epidemiology is<br />

therefore similar <strong>to</strong> that of P. graminis f. sp. tritici<br />

(Eversmeyer & Kramer, 2000). The temperature<br />

optima for urediniospore germination and infection<br />

(about 18°C) and sporulation (25°C) are<br />

lower than those of P. graminis (Singh et al.,<br />

2002). Teliospores are not readily formed, and<br />

the disease symp<strong>to</strong>ms are easily recognized by<br />

the chocolate-brown uredinial lesions on wheat<br />

leaves. There is no infection of the stem, in contrast<br />

<strong>to</strong> P. graminis. Triticale, a hybrid between<br />

wheat (Triticum) and rye (Secale), is also affected.<br />

About 50 Lr (‘leaf rust’) resistance genes are<br />

known, and wheat leaf rust is currently controlled<br />

mainly by using cultivars containing<br />

a pyramid of several of these Lr genes.<br />

22.4.2 Puccinia striiformis (stripe rust or<br />

yellow rust on wheat and barley)<br />

The alternate host of this rust species has not<br />

yet been found, and it may no longer have one.<br />

Therefore, its life cycle is probably confined <strong>to</strong><br />

the grass or cereal hosts on which uredinia and<br />

telia are produced. This rust can be distinguished<br />

from P. triticina by its yellow uredinia which are<br />

arranged in stripe-like rows following the leaf<br />

veins. There are several formae speciales, but their<br />

host ranges overlap <strong>to</strong> a greater extent than<br />

in the other cereal rusts and they are therefore<br />

less clearly delimited. The most important<br />

forms are those on wheat (f. sp. tritici) and<br />

barley (f. sp. hordei). The temperature optima of<br />

P. striiformis are the lowest of any of the common<br />

cereal rusts, with urediniospore germination<br />

and penetration around 10°C, and urediniospore<br />

production at 12 15°C (Singh et al., 2002). This<br />

species overwinters by using winter crops and<br />

volunteers as a ‘green bridge’, and can cause<br />

early epidemics in spring. Zadoks and Bouwman<br />

(1985) have estimated that a single uredinium

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