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Introduction to Fungi, Third Edition

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374 HYMENOASCOMYCETES: PYRENOMYCETES<br />

Fig12.43 Phomopsis phaseoli.(a)Surfaceviewofa<br />

conidium-producing stroma formed in agar culture.<br />

(b) Phialides producing b-conidia. (c) Ovoid a-conidium and<br />

elongated b-conidium. Both types of conidium are produced<br />

from similar phialides. (b) and (c) <strong>to</strong> same scale.<br />

like all species of Diaporthe examined <strong>to</strong> date.<br />

In culture, the mycelium consists of narrow<br />

hyphae containing 3 4 nuclei per segment, and<br />

wider ones with up <strong>to</strong> 15 nuclei per segment.<br />

Hyphae aggregate and swell <strong>to</strong> form a pseudoparenchyma<strong>to</strong>us<br />

stroma which produces both<br />

pycnidia and perithecia. In nature, the pycnidial<br />

Phomopsis state is produced earlier than the<br />

Diaporthe state which is most often seen on<br />

dead plant material. Conidial locules form in the<br />

upper region of the stroma (ec<strong>to</strong>stroma) when<br />

two opposing palisades of hyphae press against<br />

each other, accompanied by lysis of hyphae<br />

bordering the developing slit. In consequence,<br />

the slit becomes convoluted and lined<br />

by hymenium. Ec<strong>to</strong>stromatic cells proliferate <strong>to</strong><br />

form a neck, resulting in the typical shape of the<br />

pycnidium consisting of a more or less globose<br />

structure within which the lobed or folded<br />

hymenium is located, and one or several necks<br />

through which conidia are exuded (Fig. 12.43a).<br />

The conidiogenous cells are interpreted as<br />

phialides. They are awl-shaped, 20 mm long and<br />

tapering from 2 3 mm at their base <strong>to</strong> 1 mm at<br />

their apex (Fig. 12.43b). Two types of conidia are<br />

produced, often within the same conidioma<br />

(Fig. 12.43c). The ovoid a-conidia contain two<br />

lipid droplets and germinate readily in culture,<br />

whereas b-conidia are highly elongated and do<br />

not usually germinate in P. phaseoli or other<br />

Phomopsis spp. They are therefore interpreted as<br />

spermatia (Jensen, 1983).<br />

According <strong>to</strong> Uecker (1988), Phomopsis typically<br />

has a dark stroma producing a- and b-conidia,<br />

the teleomorph being Diaporthe. Unfortunately,<br />

few typical Phomopsis species exist since many<br />

produce only either a- or b-conidia, and/or<br />

lack the Diaporthe state. Further, there are few<br />

distinguishing features in conidial shape<br />

between different Phomopsis species, and species<br />

identification is based mainly on the host species<br />

with which a given strain is associated. This<br />

poses problems because the delimitations of host<br />

ranges are not precisely characterized. Thus, the<br />

taxonomy of Phomopsis is in a state of confusion,<br />

with many synonyms probably in existence.<br />

Uecker (1988) has compiled over 800 Phomopsis<br />

names in current use, but Kirk et al. (2001)<br />

have estimated that only about 100 species of<br />

Phomopsis exist. The genus is thus in urgent need<br />

of an up-<strong>to</strong>-date monographic treatment, Grove<br />

(1935) having been the last mycologist <strong>to</strong> rise<br />

<strong>to</strong> this formidable task.<br />

In D. phaseolorum, sexual reproduction is<br />

initiated by the formation of ascogonial coils in<br />

the lower region of the stroma, termed en<strong>to</strong>stroma<br />

by Jensen (1983). These coils become

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