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Introduction to Fungi, Third Edition

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MICROASCALES<br />

369<br />

Fig12.38 Scopulariopsis brevicaulis. Conidiophores terminating<br />

in conidiogenous cells (annellides) from which chains of conidia<br />

develop.The stippled areas at the tips of the annellides indicate<br />

the region of growth associated with the development of<br />

successive conidia.<br />

silage, and on keratin-rich substrates. Because<br />

of their ability <strong>to</strong> degrade keratin and grow at<br />

37°C, some Scopulariopsis spp. can be pathogenic<br />

<strong>to</strong> humans, occasionally causing deep-seated<br />

mycoses (de Hoog et al., 2000a). Not surprisingly<br />

for such versatile and adaptable fungi, they<br />

quickly develop resistance against commonly<br />

used antifungal drugs (Cuenca-Estrella et al.,<br />

2003). This feature, rather than their pathogenicity<br />

per se, seems <strong>to</strong> be the main reason why<br />

they can be troublesome in clinical situations.<br />

Several well-known Scopulariopsis states<br />

have only recently been assigned <strong>to</strong><br />

heterothallic Microascus species (Abbott & Sigler,<br />

2001). Detailed developmental studies of perithecium<br />

formation in homothallic species have<br />

been carried out by Corlett (1966). The vegetative<br />

hyphae of Microascus consist of uninucleate segments.<br />

Sexual reproduction is initiated when<br />

a coiled ascogonium forms and becomes<br />

ensheathed by hyphae arising from the ascogonial<br />

base as well as the surrounding vegetative<br />

mycelium. At this primordium stage, the outermost<br />

layer already becomes melanized, and it<br />

surrounds the developing pseudoparenchyma.<br />

The primordium enlarges by further growth of<br />

the pseudoparenchyma. The ascogonium is positioned<br />

initially in a cavity in the centre of the<br />

primordium, but soon sterile hyphae grow<br />

inwards from the surrounding pseudoparenchyma,<br />

raising the ascogonium <strong>to</strong>wards the<br />

apex of the perithecium. From the ascogonium,<br />

ascogenous hyphae grow between the sterile,<br />

inwardly radiating hyphae and form asci without<br />

croziers from their tips which have become<br />

binucleate. Simultaneously, at the tip of the<br />

perithecium pseudoparenchyma<strong>to</strong>us hyphae<br />

begin <strong>to</strong> grow inwards and then upwards,<br />

rupturing the apical wall and extending a<br />

perithecial neck with an ostiole. During maturation<br />

of the perithecium, the sterile hyphae in<br />

the perithecial centre lyse, followed by disintegration<br />

of the ascus walls, so that the mature<br />

perithecium contains an outer melanized<br />

layer surrounding a pseudoparenchyma which<br />

encloses masses of free ascospores. These ooze<br />

out through the ostiole as a tendril.<br />

12.7.2 Cera<strong>to</strong>cystis and Sphaeronaemella<br />

(Cera<strong>to</strong>cystidaceae)<br />

The most important genus of this small family<br />

is Cera<strong>to</strong>cystis (14 species) associated with living<br />

trees. Some species cause vascular wilts, perhaps<br />

the most important pathogen being C. fagacearum,<br />

the cause of oak wilt, which is especially<br />

destructive in the United States. Cera<strong>to</strong>cystis<br />

fimbriata attacks an exceptionally wide range of<br />

plants, including the European plane (Platanus<br />

acerifolia), Eucalyptus spp., mango, coffee, sweet<br />

pota<strong>to</strong> and rubber (Kile, 1993). However, infections<br />

by Cera<strong>to</strong>cystis spp. need not be destructive<br />

<strong>to</strong> the host, and some species produce benign

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