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Introduction to Fungi, Third Edition

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440 HYMENOASCOMYCETES: HELOTIALES (INOPERCULATE DISCOMYCETES)<br />

Fig15.7 Development of Pyrenopeziza brassicae ascocarps. (a) Immature apothecium,10 days old on oilseed rape.The apical<br />

pore has just formed. (b) 14-day-old apothecium on oilseed rape.The opening is widening due <strong>to</strong> the expansion of the basal disc.<br />

(c) Mature apothecium 46 days after inoculation on<strong>to</strong> agar. Reprinted from Gilles et al. (2001), with permission from Elsevier.<br />

Images kindly provided by N. Evans.<br />

(Salamati et al., 2000). The mating type idiomorphs<br />

have been characterized for all species<br />

except Tapesia acuformis (Foster & Fitt, 2004), and<br />

they are of the usual heterothallic/bipolar type.<br />

The biology of the two sister species Tapesia<br />

yallundae on wheat and T. acuformis on rye<br />

(formerly called T. yallundae W and R pathotypes,<br />

respectively) is very similar and has been<br />

reviewed by Fitt et al. (1988) and Lucas et al.<br />

(2000). Eyespot is a major disease in winter<br />

cereals growing in cool climates. Infection is<br />

probably mainly by the needle-shaped conidia<br />

which are formed on overwintered stubble and<br />

spread by rain splash. However, ascospores<br />

released from apothecia (Fig. 15.8a) in early<br />

spring are also infectious. If a spore lands on<br />

the coleoptile of a host plant, it germinates and<br />

produces an aggregate of hyphae termed an<br />

infection plaque (Fig. 15.8b). Numerous melanized<br />

appressoria are formed at the interface<br />

of this structure with the host epidermis, so that<br />

infection of susceptible hosts occurs at several<br />

points (Fig. 15.8c). Penetration is probably<br />

mediated by a combination of turgor pressure<br />

and hydrolytic enzymes. The typical eyespot<br />

(Fig. 15.8d) develops as a greyish-brown lesion<br />

around clusters of infection plaques which may<br />

be visible as the ‘pupil’ of the eyespot. Detailed<br />

studies of infection mechanisms have been<br />

published by Daniels et al. (1991, 1995). The presence<br />

of eyespots at the haulm bases renders the<br />

cereal shoots prone <strong>to</strong> collapsing. Further infections<br />

can affect the vascular system, resulting<br />

in poorly developed ‘whiteheads’ containing<br />

inferior grain.<br />

Resistance breeding seems <strong>to</strong> be a promising<br />

strategy for the control of eyespot in cereals<br />

(Lucas et al., 2000). Chemical control is also practised,<br />

but Tapesia spp. have developed resistance<br />

against several types of fungicide (Leroux &<br />

Gredt, 1997).<br />

15.4 Rhytismataceae<br />

The taxonomy of this family is still in a state of<br />

flux (Gernandt et al., 2001). It is sometimes given<br />

ordinal status (Rhytismatales or Phacidiales).<br />

The apothecia are immersed in host tissue or<br />

embedded in a flat stroma. Individual apothecia<br />

become evident when the upper surface breaks<br />

open <strong>to</strong> reveal the hymenium. There are<br />

219 species in this group at present (Kirk et al.,<br />

2001). Most of them are associated with broadleaved<br />

trees or conifers (Cannon & Minter, 1986;<br />

Johns<strong>to</strong>n, 1997). Particularly difficult genera

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