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Introduction to Fungi, Third Edition

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DOTHIDEALES<br />

483<br />

Fig17.20 Wheat leaf blotch symp<strong>to</strong>ms caused by<br />

Mycosphaerella graminicola. (a) Infected leaf showing a necrotic<br />

lesion, in the centre of which pycnidia of Sep<strong>to</strong>ria tritici have<br />

formed (arrows). (b) Septate conidia of S. tritici,morethan<br />

10 times longer than wide.<br />

its pseudothecia on overwintered stalks and<br />

leaves of numerous monocotyledons and dicotyledons<br />

in subarctic and subalpine regions, and a<br />

period of cold is required for ascocarp initiation<br />

in culture (Barr, 1958; Corlett, 1991). In contrast,<br />

C. herbarum is ubiqui<strong>to</strong>us and common in<br />

temperate regions on senescent and dead plant<br />

material, and in soil. Conidia of this and other<br />

Cladosporium spp. are the most abundant component<br />

of the fungal air spora (Gregory, 1973), and<br />

they are probably the most frequent contaminant<br />

of foodstuffs, textiles and paintwork.<br />

They also frequently contaminate cultures of<br />

other fungi in the labora<strong>to</strong>ry. The conidia of<br />

C. herbarum and other common moulds such as<br />

Alternaria alternata and Aspergillus fumigatus are<br />

associated with severe asthma (Zureik et al.,<br />

2002). Over 30 antigens causing mould allergy<br />

have been described from C. herbarum, and most<br />

of them are secre<strong>to</strong>ry or cy<strong>to</strong>plasmic glycoproteins,<br />

often representing common enzymes such<br />

as enolase or aldehyde dehydrogenase<br />

(Breitenbach & Simon-Nobbe, 2002).<br />

Colonies of C. herbarum are dull olive green<br />

<strong>to</strong> black in colour, and appear as a network<br />

of hyphae or a plate-like mass (stroma) of<br />

tightly packed, dark, thick-walled cells (McKemy<br />

& Morgan-Jones, 1991). The conidiophores are<br />

branched or unbranched and conidiogenesis is<br />

holoblastic (Fig. 17.23a). The tip of the conidiophore<br />

bulges out <strong>to</strong> form the first conidium and<br />

it is presumed that all the wall layers of the apex<br />

are involved (Hashmi et al., 1973). The first<br />

conidium buds <strong>to</strong> form a further conidium and<br />

this process continues so that a chain of conidia<br />

develops in acropetal succession, the youngest<br />

conidium at the end of the chain. Most conidia<br />

have a scar (hilum) at each end, but occasionally<br />

a conidium may form two daughter conidia<br />

at its tip so that, as further conidial development<br />

proceeds, a branched chain develops (see<br />

Fig. 17.23a). Such branch-point conidia have been<br />

termed ramoconidia (Lat. ramus ¼ branch) and<br />

are marked by having a single scar at the base<br />

and two scars at the apex. The conidia of<br />

C. herbarum have dark (melanized) walls which<br />

are slightly roughened. They may remain unicellular<br />

or develop 1 3 transverse septa. Another<br />

common species of Cladosporium is C. cladosporioides,<br />

which has smooth conidium walls.<br />

Cladosporium fulvum (also called Fulvia fulva<br />

or Mycovellosiella fulva) is probably not closely<br />

related <strong>to</strong> other Cladosporium spp. No sexual state<br />

has been reported, but other Mycovellosiella<br />

spp., like Cladosporium spp., belong <strong>to</strong><br />

Mycosphaerella (Crous et al., 2001). In contrast <strong>to</strong><br />

most other plant-pathogenic members of the<br />

Loculoascomycetes, C. fulvum is biotrophic. It is<br />

a pathogen of <strong>to</strong>ma<strong>to</strong> plants, especially in<br />

greenhouses. Conidia infect their host through<br />

s<strong>to</strong>mata, and hyphae spread in the leaf apoplast<br />

in close contact with mesophyll cells, but without<br />

producing haus<strong>to</strong>ria. Sucrose, the major<br />

plant transport sugar, is hydrolysed and taken<br />

up, being converted <strong>to</strong> manni<strong>to</strong>l by the fungus<br />

(Joosten et al., 1990). Conspicuous yellow

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