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Introduction to Fungi, Third Edition

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484 LOCULOASCOMYCETES<br />

incompatible interactions results if the product<br />

of the Avr gene is recognized by the host plant.<br />

Several Avr genes have been characterized, and<br />

their products are usually small proteins<br />

secreted by C. fulvum in<strong>to</strong> the apoplast. Many of<br />

the corresponding Cf genes of <strong>to</strong>ma<strong>to</strong> are also<br />

known; they encode proteins anchored in the<br />

plasma membrane of <strong>to</strong>ma<strong>to</strong> cells, with large<br />

extracellular domains. The examination of the<br />

products of matching avirulence and resistance<br />

genes should provide an opportunity <strong>to</strong> examine<br />

their interactions, and thus the molecular basis<br />

of recognition events involved in specific resistance<br />

(Rivas & Thomas, 2005). This work is still<br />

ongoing.<br />

Fig17.21 Cercospora beticola.Conidiophores and conidia from<br />

sugar beet seed.<br />

(chlorotic) leaf areas are produced as a result of<br />

such systemic infections (Fig. 17.24), and eventually<br />

conidiophores are emitted through<br />

s<strong>to</strong>mata especially on the lower leaf surface,<br />

forming a lawn of spores resembling powdery<br />

mildews but being light brown in colour.<br />

The interaction between C. fulvum and its<br />

<strong>to</strong>ma<strong>to</strong> host is governed by a classical genefor-gene<br />

relationship based on dominant host<br />

resistance genes (Cf genes, for C. fulvum) and<br />

dominant avirulence (Avr) genes in C. fulvum,<br />

i.e. virulence is a recessive trait (Joosten et al.,<br />

1997). The hypersensitive response of<br />

17.3.5 Aureobasidium and black yeasts<br />

Aureobasidium pullulans is a ubiqui<strong>to</strong>us saprotroph<br />

whose main habitats are the phylloplane<br />

and other surfaces of living and senescent plants,<br />

but it can also occur as a symp<strong>to</strong>mless endophyte.<br />

It grows in soil, but is often not recorded<br />

because it is temperature-sensitive and is not<br />

seen on soil plates prepared with warm agar. It<br />

has been isolated from fresh water, estuarine<br />

and marine sediments, sea water, sewage and<br />

other liquid waste (Domsch et al., 1980). Its<br />

teleomorph is Discosphaerina fulvida, a relative of<br />

Mycosphaerella (Yurlova et al., 1999). The fungus<br />

can be readily isolated from leaf washings. It is<br />

pleomorphic, and in culture it forms a rapidly<br />

growing mycelium with wide, septate hyphae<br />

from which intercalary and occasionally terminal<br />

cells give rise <strong>to</strong> single or clustered hyaline<br />

blas<strong>to</strong>conidia by a process of budding. Budding is<br />

associated with local lysis of the wall of the<br />

conidiogenous cell, the inner wall of which then<br />

balloons out and forms the wall of the conidium<br />

(Ramos et al., 1975). Repeated conidium development<br />

from the same or closely adjacent conidiogenous<br />

loci results in the formation of<br />

slimy clusters of hyaline conidia (Fig. 17.25a).<br />

Intercalary cells may enlarge and develop<br />

thicker, dark, melanized walls <strong>to</strong> become<br />

chlamydospores (Fig. 17.25b). Conidia bud in a<br />

yeast-like manner when the fungus is grown<br />

in liquid culture with high inoculum densities.<br />

The yeast cells can continue <strong>to</strong> bud

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