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Introduction to Fungi, Third Edition

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598 HETEROBASIDIOMYCETES<br />

(Uetake et al., 1992), and the same pro<strong>to</strong>corm,<br />

mature root or even individual cell can be<br />

colonized simultaneously by different fungi.<br />

The unstable nature of the orchid mycorrhiza<br />

is indicated by the quick and repeated cycle of<br />

pelo<strong>to</strong>n formation and degradation, and the<br />

several different outcomes of the orchid fungus<br />

interaction observed under labora<strong>to</strong>ry conditions<br />

(Fig. 21.2d). A balanced mycorrhizal symbiosis<br />

will develop only in a proportion of<br />

pro<strong>to</strong>corms, whereas other seeds of the same<br />

orchid species may be parasitized and killed by<br />

the fungus, or simply resist infection and stall in<br />

their development (Hadley, 1970; Smreciu &<br />

Currah, 1989; Beyrle et al., 1995). There is also<br />

vevidence of a succession of mycorrhizal fungi<br />

during the development of an orchid in nature,<br />

and the mycorrhizal fungi isolated from adult<br />

plants may not support pro<strong>to</strong>corm growth and<br />

vice versa (Xu & Mu, 1990; Zelmer et al., 1996).<br />

Mature orchids may be associated with<br />

Rhizoc<strong>to</strong>nia spp. and/or a range of other Basidiomycota,<br />

including saprotrophic (e.g. Mycena),<br />

necrotrophic (e.g. Armillaria; see p. 546) or<br />

ec<strong>to</strong>mycorrhizal species related <strong>to</strong> Russula and<br />

Thelephora (Rasmussen, 2002). Ec<strong>to</strong>mycorrhizal<br />

fungi transport carbohydrates from their tree<br />

host <strong>to</strong> the orchid, thus allowing green orchids<br />

<strong>to</strong> grow even in densely shaded woodland<br />

conditions (Taylor & Bruns, 1997; McKendrick<br />

et al., 2000; Bidar<strong>to</strong>ndo et al., 2004). Interestingly,<br />

these fungi form typical ec<strong>to</strong>mycorrhiza with<br />

their tree hosts but pelo<strong>to</strong>ns in infected orchid<br />

roots (Zelmer & Currah, 1995). The orchid therefore<br />

calls the shots in its symbiosis with<br />

basidiomycetes, and pelo<strong>to</strong>n formation and<br />

degradation is traditionally interpreted as a<br />

balanced defence reaction of the orchid against<br />

invasion attempts by the fungus.<br />

21.3 Dacrymycetales<br />

This order is characterized by forked (furcate)<br />

basidia (Figs. 21.4 and 21.5), which are found in<br />

all species except Dacrymyces unisporus. The fruit<br />

bodies are coloured yellow or orange due <strong>to</strong> the<br />

presence of a wide range of carotenoids (for<br />

references, see Gill & Steglich, 1987). Fruit bodies<br />

are gelatinous and show a striking diversity of<br />

forms as exemplified by the cushion-like basidiocarps<br />

of Dacrymyces stillatus (Fig. 21.3) and the<br />

clavarioid ones of Calocera viscosa (Plate 11g).<br />

Not much is known about the life cycle of<br />

the Dacrymycetales, but it is presumed that<br />

the usual basidiomycete pattern of alternating<br />

mono- and dikaryotic stages operates. There are<br />

no clamp connections. The dolipore-type septa<br />

Fig 21.3 Fruit bodies of Dacrymycetales. (a) Basidial<br />

cushions of Dacrymyces stillatus on rotting wood.<br />

(b) Basidiocarps of Calocera cornea.

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