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Introduction to Fungi, Third Edition

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472 LOCULOASCOMYCETES<br />

Table 17.2. Selected plant-pathogenic species of Alternaria (from Holliday, 1998).<br />

Species of Alternaria Disease Comments<br />

‘A. alternata’<br />

Widerangeofhostsanddiseases<br />

‘A. alternata’f. sp. lycopersici Toma<strong>to</strong> stem canker<br />

A. brassicae Brassica greyleaf spot Seed-borne<br />

A. brassicicola Brassica blackleaf spot Seed-borne<br />

A. carthami Safflower leaf spot Seed-borne<br />

A. dauci Carrot leaf blight Seed-borne<br />

A. dianthi Carnationleaf blight Seed-borne<br />

A. linicola Linseed seedling blight Seed-borne<br />

A. macrospora Cot<strong>to</strong>nleaf spot Seed-borne<br />

A. mali Apple core rot<br />

A. porri Onion purple blotch Seed-borne<br />

A. radicina Carrot black rot; also infects celery and parsnip<br />

A. solani Pota<strong>to</strong> early blight and tuber rot<br />

Toma<strong>to</strong> early blight and fruit rot<br />

Curvularia (Fig. 17.14). The revised genus<br />

Helminthosporium is now rather small, with<br />

about 20 species and H. velutinum as the typespecies<br />

(Fig. 17.12). It has affinities <strong>to</strong><br />

Lep<strong>to</strong>sphaeria (Olivier et al., 2000). Drechslera, in<br />

contrast, is the anamorphic state of Pyrenophora<br />

(Fig. 17.16; see p. 477). All these anamorphic<br />

forms produce pigmented (melanized) spores<br />

with only transverse septa, but they differ in<br />

their pattern of conidiogenesis and in the<br />

ultrastructural appearance of their cell walls.<br />

Sporogenesis by Helminthosporium is tretic, i.e.<br />

porospores are formed as described for<br />

Stemphylium by the digestion of a small hole<br />

in<strong>to</strong> the conidiophore cell wall, followed by an<br />

extension of the inner wall <strong>to</strong> form the conidium<br />

which then lays down its outer wall (Fig. 17.12c).<br />

In contrast <strong>to</strong> Pleospora, each site can produce<br />

only one conidium, and conidia can develop<br />

apically or laterally on the conidiophore. If the<br />

conidium is produced apically, the conidiophore<br />

cannot grow further. The sequence of conidium<br />

development in Bipolaris and Curvularia is not<br />

of this type. In Bipolaris, the first conidium<br />

always develops apically, and subsequent<br />

conidia are formed either by growing through<br />

the scar left by the first conidium, or by the<br />

conidiophore growing past the first conidium <strong>to</strong><br />

form a new apex producing the second conidium<br />

(B. sorokiniana; Fig. 17.13b). Bipolaris is so named<br />

because the conidia germinate by emitting two<br />

germ tubes, one at either end (Fig. 17.13d), and<br />

these grow as extensions of the long axis of the<br />

spore. This is in contrast <strong>to</strong> Helminthosporium<br />

conidia in which each cell is principally capable<br />

of germination, and the germ tubes grow<br />

perpendicular <strong>to</strong> the long axis of the spore.<br />

In Curvularia, the conidia are curved because<br />

of an unevenly swollen central cell. The end cells<br />

are usually less strongly pigmented than the<br />

central cell (Figs. 17.14b,c). The first conidium<br />

also develops at the apex of the conidiophore as<br />

a poroconidium extending through a tiny pore,<br />

and then the conidiophore develops a new<br />

subterminal growing point from which a<br />

second conidium initial arises. The process is<br />

repeated so that a succession of new apices, each<br />

terminated by a conidium, is formed (Fig. 17.14a).<br />

The term sympodula has been applied <strong>to</strong> such<br />

a conidiophore producing conidia sympodially<br />

(Kendrick & Cole, 1968). In some Curvularia spp.,<br />

such as C. cymbopogonis (Fig. 17.14c), the base of<br />

the conidium bears a protuberant hilum.<br />

A feature of the conidia of Drechslera,<br />

Helminthosporium and Bipolaris, but not<br />

Curvularia, is that they are dis<strong>to</strong>septate<br />

(Luttrell, 1963). This means that the wall separating<br />

adjacent conidial segments is visibly

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