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Introduction to Fungi, Third Edition

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620 UREDINIOMYCETES: UREDINALES (RUST FUNGI)<br />

s<strong>to</strong>matal surface acting as the thigmotropic<br />

signal (Rubiales & Niks, 1996). There is also<br />

adult-plant resistance, usually an interaction<br />

between various genes which retard the progress<br />

of infection <strong>to</strong> such a degree that the losses are<br />

reduced <strong>to</strong> economically acceptable levels.<br />

22.3 Puccinia graminis, the cause of<br />

black stem rust<br />

A connection between the barberry bush (Berberis<br />

vulgaris) and rust disease on wheat and other<br />

cereals had been suspected for centuries by<br />

farmers who noticed the frequent occurrence of<br />

crop damage around or downwind from barberry<br />

bushes. Further, an al<strong>to</strong>gether different-looking<br />

fungus known as Aecidium berberidis was known<br />

<strong>to</strong> infect barberry leaves. The formal proof that<br />

the barberry and wheat pathogens were different<br />

stages of the same species was made by An<strong>to</strong>n de<br />

Bary who, in 1865, performed cross-inoculation<br />

experiments <strong>to</strong> show that basidiospores derived<br />

from germinating teliospores from wheat were<br />

able <strong>to</strong> cause spermogonial and aecial infections<br />

on the barberry. This s<strong>to</strong>ry has been recounted<br />

many times, but nobody has <strong>to</strong>ld it better than<br />

Large (1940).<br />

Black stem rust affects especially wheat but<br />

also most other cereals and a range of wild<br />

grasses. Crop losses can be severe as a reduction<br />

in quantity as well as quality of the grain yield.<br />

The host epidermis can become ruptured over<br />

much of its surface in severe infections, thereby<br />

debilitating the plants (see Fig. 22.8d). Serious<br />

epidemics have occurred in the past, e.g. in the<br />

USA in 1904 and especially during the war year<br />

1916. In fact, losses were so severe (up <strong>to</strong> 50% in<br />

the Great Plains; Eversmeyer & Kramer, 2000)<br />

that P. graminis was seriously considered and<br />

developed as a biological weapon by the US<br />

Government during the 1960s (Line & Griffith,<br />

2001). On susceptible wheat cultivars and without<br />

chemical protection, P. graminis can cause<br />

<strong>to</strong>tal crop failure.<br />

Although it is an ecologically obligate<br />

biotroph, Puccinia graminis is able <strong>to</strong> infect an<br />

as<strong>to</strong>nishingly wide range of grass and cereal<br />

hosts. Gäumann (1959) listed 365 host species in<br />

Fig 22.8 Puccinia graminis. (a) Spermogonial pustules on the upper surface of a leaf of Berberis vulgaris.Notethedropsofnectar.<br />

(b) Aecia on the underside of a Berberis leaf.The outer frilly layer is the white peridium, within which is a mass of orange-coloured<br />

aeciospores. (c) Wheat leaf showing uredinia which appear as reddish-brown powdery masses. (d) Wheat straw showing telia as<br />

black raised pustules.

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