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Introduction to Fungi, Third Edition

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BLUMERIA GRAMINIS<br />

395<br />

Fig13.3 An example of the use of scanning electron microscopy in fungal taxonomy, separating Blumeria graminis (a,b) from other<br />

members of the Erysiphales (c,d). (a) Whole conidium of B. graminis showing the spiny surface of the wall. (b) The‘annular’ septum<br />

of the conidium.The actual septum is raised and warty, but it is surrounded by a depressed annulus. (c) Conidium of Golovinomyces<br />

(Erysiphe) cynoglossi showing a slightly roughened wall. (d) Conidium of Neoerysiphe (Erysiphe) galeopsidis with its striate conidial wall.<br />

Micrographs kindly provided by R.T. A.Cook. Reprinted from Cook et al. (1997), British Crown copyright1997.<br />

The genus Blumeria contains only one species,<br />

B. graminis, which is separable in<strong>to</strong> several<br />

formae speciales distinguishable by their different<br />

grass (Agropyron, Bromus, Poa) or cereal hosts<br />

(barley, oat, rye, wheat). Thus, B. graminis f. sp.<br />

tritici infects wheat (Triticum) but not barley,<br />

whilst B. graminis f. sp. hordei infects barley<br />

(Hordeum) but not wheat. Blumeria graminis is heterothallic<br />

and hybrids between certain formae<br />

speciales may arise, especially if the hosts themselves<br />

can hybridize. For example, it has been<br />

shown that hybridization between B. graminis ff.<br />

spp. agropyri, tritici and secalis can occur, resulting<br />

in viable ascospores. The three host genera<br />

Agropyron (couch grass), Triticum (wheat) and<br />

Secale (rye) can also hybridize (Hiura, 1978).<br />

Recently, it has become obvious that the definition<br />

of formae speciales is less clear-cut than previously<br />

thought. Extensive overlaps in host spectra<br />

occur especially on wild hosts in the Middle East<br />

where Hordeum spontaneum, the presumed ances<strong>to</strong>r<br />

species of cultivated barley, is endemic and<br />

where B. graminis probably originated (Clarke &<br />

Akhkha, 2002; Wyand & Brown, 2003). Further,<br />

within a given forma specialis, numerous races<br />

can be distinguished by specific features such<br />

as resistance <strong>to</strong> fungicides or ability <strong>to</strong> infect<br />

a given host cultivar.<br />

13.3.1 The infection process<br />

A conidium of B. graminis alighting on the surface<br />

of a grass or cereal leaf will initially come <strong>to</strong> rest<br />

by the tips of its spines. Within seconds of contact<br />

with hydrophobic (but not with hydrophilic)<br />

surfaces, an extracellular matrix is released by<br />

those spines <strong>to</strong>uching the surface. The matrix<br />

contains proteins, including enzymes such as<br />

cutinases and non-specific esterases, and it may<br />

serve in the initial attachment of the conidium <strong>to</strong><br />

the surface (Carver et al., 1999; Nielsen et al., 2000).

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