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Introduction to Fungi, Third Edition

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540 HOMOBASIDIOMYCETES<br />

is used in attempts <strong>to</strong> wean alcoholics from their<br />

addiction, although it is different chemically.<br />

Interference competition involving Coprinus<br />

Many species of Coprinus are coprophilous, typically<br />

with their basidiocarps appearing relatively<br />

late in the succession of fungi on dung (Dix &<br />

Webster, 1995). Certain species are known <strong>to</strong><br />

suppress the fruiting of other fungi. A good<br />

example of this phenomenon is C. heptemerus,<br />

which inhibits the fruiting of many species in<br />

nature on rabbit dung and in culture. Its effect<br />

on the sensitive ascomycete Ascobolus crenulatus<br />

was tracked down <strong>to</strong> the moment when the<br />

hyphae of the two fungi make contact. Within<br />

minutes, the hyphal segment of Ascobolus<br />

<strong>to</strong>uched by a hyphal tip of Coprinus is killed.<br />

There is a rapid loss of turgor, shown by the<br />

bulging of the septa of adjacent cells in<strong>to</strong> the<br />

affected cell, which also loses the ability <strong>to</strong><br />

undergo plasmolysis, whilst adjacent cells readily<br />

plasmolyse when bathed in hyper<strong>to</strong>nic fluid.<br />

This form of competition has been termed<br />

hyphal interference or interference competition.<br />

It occurs in a range of genera of coprophilous<br />

basidiomycetes, but is not confined <strong>to</strong> this ecological<br />

group, having been demonstrated <strong>to</strong> be<br />

effective also in lignicolous fungi (Ikediugwu &<br />

Webster, 1970a,b; Ikediugwu et al., 1970).<br />

Cell damage is very similar <strong>to</strong> the effects seen<br />

when self- and non-self hyphae of the same<br />

species confront each other. An oxidative burst is<br />

stimulated and hydrogen peroxide accumulates<br />

in cells of the sensitive partner. This is interpreted<br />

as a defence reaction (Silar, 2005).<br />

19.4.3 Amanitaceae<br />

Amanita (c. 500 spp.)<br />

This is a large and important genus whose<br />

species form sheathing mycorrhiza with trees.<br />

Amanita muscaria (‘fly agaric’; Plate 9a) is often<br />

associated with birch (Betula) but also grows in<br />

mycorrhizal association with Abies, Pinus, Picea,<br />

Quercus and other hosts. As is well known,<br />

the basidiocarps of some species are poisonous,<br />

especially those of A. phalloides (death cap),<br />

A. virosa (destroying angel), A. pantherina (panther<br />

cap) and A. verna, whilst those of A. muscaria are<br />

more hallucinogenic than poisonous. There are<br />

also species whose basidiocarps are excellent <strong>to</strong><br />

eat, most notably A. caesarea (Caesar’s mushroom;<br />

Plate 9b), which has been hunted enthusiastically<br />

in Southern European countries since Roman<br />

times. Emperor Claudius was an early connoisseur<br />

of A. caesarea and may have paid for his<br />

mycophagy with his life, probably falling victim<br />

<strong>to</strong> a poisoned mushroom dish manipulated by<br />

his wife Agrippina in AD 54 (Ramsbot<strong>to</strong>m, 1953).<br />

Other edible species are A. rubescens (blusher),<br />

A. vaginata (grisette) and A. fulva (tawny grisette).<br />

In view of the possible confusion between edible<br />

and poisonous species, it is obviously best <strong>to</strong><br />

avoid eating basidiocarps of any whose identity<br />

is uncertain.<br />

The characteristic features of Amanita include<br />

a white spore print and the presence of a volva,<br />

i.e. the <strong>to</strong>rn remnants of a universal veil. The<br />

volva persists as a cup at the base of the stipe and<br />

broken volva fragments may also adhere <strong>to</strong> the<br />

cap, as seen as the white scales on the red caps of<br />

A. muscaria (Plate 9a). Most species also have<br />

a ring (annulus) on the stem, the remnant of the<br />

partial veil which protected the gills during fruit<br />

body development, but there is no ring in<br />

some species formerly classified in Amani<strong>to</strong>psis<br />

(e.g. A. vaginata and A. fulva).<br />

Amanita poisoning<br />

Symp<strong>to</strong>ms after ingestion of fruit bodies of<br />

A. phalloides follow a characteristic time course<br />

over a period of 7 days (Faulstich & Zilker, 1994).<br />

After a mushroom meal and symp<strong>to</strong>m-free<br />

interval (day 1), there is a period of emesis<br />

(vomiting), abdominal cramps and diarrhoea<br />

(day 2), followed by a period of remission (day<br />

3) which is treacherous because it lures many<br />

patients in<strong>to</strong> believing that they have overcome<br />

the poisoning. Meanwhile, severe liver damage is<br />

ongoing and symp<strong>to</strong>ms resume with a vengeance<br />

with gastrointestinal bleeding (day 4), hepatic<br />

encephalopathy (brain damage, day 5), kidney<br />

failure (day 6) and death (day 7).<br />

Amanita poisoning is caused by two <strong>to</strong>xins,<br />

namely the ama<strong>to</strong>xin a-amanitin and the<br />

phallo<strong>to</strong>xin phalloidin (Bresinsky & Besl, 1990;<br />

Wieland & Faulstich, 1991; Chil<strong>to</strong>n, 1994,<br />

Wieland, 1996). Both are bicyclic oligopeptides

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