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Introduction to Fungi, Third Edition

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500 BASIDIOMYCOTA<br />

Fig18.11 Diagrammatic representation of the<br />

sequence of events associated with the formation<br />

of a clamp connection in the dikaryotic hypha of a<br />

basidiomycete. (a) Terminal segment of a hypha<br />

with two genetically dissimilar nuclei. (b) A lateral<br />

bulge in the hypha appears near the paired nuclei.<br />

The leading nucleus moves in<strong>to</strong> the bulge. (c) The<br />

two nuclei undergo conjugate (i.e. simultaneous)<br />

nuclear division. Mi<strong>to</strong>sis of the leading nucleus<br />

occurs within the bulge. (d) The lateral bulge has<br />

developed in<strong>to</strong> a backwardly directed branch or<br />

hook with one daughter of the leading nucleus at<br />

its tip. One of the daughter subterminal nuclei<br />

moves forward.Transverse septa have developed<br />

simultaneously in the main hypha and at the base of<br />

the hook. (e) The tip of the hook has fused with the<br />

wall of the main hypha. Its nucleus has moved in<strong>to</strong><br />

the main hypha and taken up position behind the<br />

daughter of the subterminal nucleus. (f) The pairs<br />

of nuclei move away from the transverse septum in<br />

the main hypha, the terminal pair moving nearer<br />

the hyphal tip and the subterminal pair distally.<br />

Note that both segments contain dissimilar nuclei<br />

but that their arrangement has been reversed.<br />

nuclei, there would be a tendency for dikaryotic<br />

mycelia <strong>to</strong> break down in<strong>to</strong> homokaryotic<br />

segments. Whilst it is reasonable <strong>to</strong> infer that<br />

mycelia with clamps at the septa are dikaryotic,<br />

the converse is not true because there are<br />

numerous basidiomycetes in which the dikaryotic<br />

mycelium does not bear clamps, and this is<br />

especially true of hyphae making up basidiocarps.<br />

For a fuller discussion of dikaryon<br />

formation see Cassel<strong>to</strong>n and Economou (1985).<br />

18.7.5 Aggregates of vegetative hyphae<br />

Basidiomycete hyphae can aggregate <strong>to</strong> form<br />

complex vegetative structures such as hyphal<br />

strands, mycelial cords, mycelial sheets and<br />

rhizomorphs (Butler, 1966; Watkinson, 1979;<br />

Rayner et al., 1985). These often grow more<br />

rapidly than individual hyphae, connect food<br />

bases such as litter fragments in soil, fallen logs<br />

and tree stumps, and are capable of rapid twoway<br />

conduction of water and nutrients.<br />

Mycelial cords<br />

Mycelial cords have been defined by Boddy (1993)<br />

as aggregations of predominantly parallel,<br />

longitudinally aligned hyphae (see Fig. 1.13).<br />

They are especially common in wood-decaying<br />

basidiomycetes but are also involved in the<br />

underground spread of basidiomycetes forming<br />

ec<strong>to</strong>mycorrhiza. Wood-decaying mycelial cord<br />

formers are a very successful ecological group,<br />

using their cords in a variety of strategies <strong>to</strong><br />

secure resources, e.g. by displacing other fungi<br />

from food bases or exploring new resources in<br />

the soil. The cords show relatively little differentiation<br />

in structure and pigmentation.<br />

Examples of fungi with mycelial cords are the<br />

stinkhorn (Phallus impudicus) and the agaric<br />

Megacollybia platyphylla. Their white cords can<br />

extend for many metres in the soil and can be<br />

traced back from the fruit bodies <strong>to</strong> tree stumps<br />

or other food bases if highly motivated students<br />

and digging <strong>to</strong>ols are available (Fig. 18.13a).<br />

Rhizomorphs<br />

The term rhizomorph refers <strong>to</strong> the root-like form<br />

of these mycelial aggregates which are more<br />

highly differentiated than mycelial cords and are<br />

often pigmented brown or black due <strong>to</strong> the<br />

presence of melanin in the walls of small,

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