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Growth, Differentiation and Sexuality

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2003; Kües et al. 2004) are the first fruiting bodyspecific<br />

structures, <strong>and</strong> this stage can be seen by the<br />

naked eye using the vital stain Janus green (Sánchez<br />

<strong>and</strong> Moore 1999; Sánchez et al. 2004). Within the<br />

core of the initial, differentiation of cells occurs<br />

(Moore 1998; Kües 2000; Kües et al. 2004). The<br />

lower part will develop into the stipe, while the<br />

capwillbeformedfromtheupperpart.Withinthe<br />

cap, different tissues develop, which are not formed<br />

from meristems, as in plants, but result from the interaction<br />

of individual hyphae. By growing at their<br />

apices, they form an interwoven structure called<br />

a plectenchyma. The outer part of the cap is called<br />

the veil. In the inner part, the pileus trama <strong>and</strong> gills<br />

Fig. 19.1. Within the primordium of C. cinera,allparts<strong>and</strong><br />

tissues of the fruiting body can already be distinguished.<br />

The mature fruiting body results from intercalary cell expansion<br />

(courtesy of M. Navarro-González)<br />

Fruiting in Basidiomycetes 395<br />

(or pores) with a hymenium can be distinguished<br />

(Fig. 19.1). In the hymenium, different cell types are<br />

formed, among which the basidia. In the basidia,<br />

karyogamy <strong>and</strong> meioses take place, ultimately resulting<br />

in basidiospores.<br />

That development of fruiting bodies is complex<br />

is also exemplified by the fact that formation of the<br />

different tissues overlaps in time. Moreover, cells<br />

in the developing mushroom differ in diameter,<br />

length, the number of septa, nuclei <strong>and</strong> vacuoles as<br />

well as the molecular composition (e.g. the content<br />

of reserve carbohydrate; Moore 1998). The different<br />

cell types are not only the result of localized<br />

growth but also apoptosis is involved (Umar <strong>and</strong><br />

van Griensven 1997; Chap. 9, this volume). Fruiting<br />

bodies of other basidiomycetes, such as those of<br />

S. commune, follow a completely different morphogenetic<br />

pathway (Fig. 19.2). These fruiting bodies<br />

result from indeterminate growth of fruiting-body<br />

primorida. Expansion of the cup-shaped primordia<br />

is not the result of intercalary growth but is<br />

due to continued apical growth <strong>and</strong> differentiation<br />

of hyphae in the primordium (Wessels 1993a).<br />

The variety in developmental programmes makes<br />

it even more difficult to underst<strong>and</strong> processes involved<br />

in fruiting-body formation in the homoba-<br />

Fig. 19.2.A–D Fruit-body development of S. commune (extracted<br />

from Wessels 1996). A Cryo-scanning electron micrograph<br />

of an early stage of fruiting-body formation. Hyphae<br />

which extend at their tips aggregate <strong>and</strong> grow upwards.<br />

B As in A but at a later stage of development. C Macrograph<br />

of fruiting bodies at the cup stage. Note the formation of<br />

the split gills. D Full-grown fruiting bodies. Bars represent<br />

1 mm (A), 0.1 mm (B)<strong>and</strong>10mm (C, D)

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