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Introduction to Fungi, Third Edition

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496 BASIDIOMYCOTA<br />

mushroom Agaricus campestris produced 1.8 10 9<br />

spores over 2 days at an average rate of<br />

40 million h 1 . Estimates for some other basidiomycetes<br />

are given in Table 18.1. The mycelia of<br />

all these fungi are perennial and an individual<br />

mycelium may produce numerous basidiocarps<br />

over a period of many years. It is therefore clear<br />

that an individual basidiospore has an infinitesimal<br />

chance of successfully establishing a<br />

fruiting mycelium.<br />

Fig18.8 Representation of events associated<br />

with ballis<strong>to</strong>spore discharge. (a) Ballis<strong>to</strong>spore<br />

attached <strong>to</strong> its sterigma before drop<br />

formation.The closed circle within the spore<br />

indicates the centre of mass of the spore.<br />

(b) Buller’s drop appears at the hilar appendix.<br />

The adaxial drop emerges on the spore wall<br />

above it and extends downwards as it<br />

increases in size.The centre of mass of spore<br />

plus drop moves <strong>to</strong> a position nearer the hilar<br />

appendix. (c) Contact between the two drops<br />

is followed by immediate coalescence and the<br />

combined mass of liquid moves rapidly up the<br />

adaxial face of the spore away from the hilar<br />

appendix.The centre of mass moves very<br />

rapidly in the direction of the thin arrow and<br />

the spore drop system gains kinetic energy<br />

and momentum in the same direction,<br />

simultaneously exerting an opposite force F<br />

on the sterigma at the hilum (thick arrow).<br />

Some angular momentum is also exerted on<br />

the spore, related <strong>to</strong> the distance a between<br />

the hilum and the hilar appendix. Reprinted<br />

from Webster et al. (1988), with permission<br />

from Elsevier.<br />

18.7 Basidiospore germination and<br />

hyphal growth<br />

18.7.1 Germination<br />

Basidiospores may remain dormant and retain<br />

viability for several months or even for a few<br />

years if conditions are unsuitable for germination.<br />

Dormancy is frequently exogenous, i.e. the<br />

spores require some external chemical or physical<br />

stimulus before germination can occur.<br />

Germination may be direct by production of a<br />

germ tube, by repetition (i.e. the formation of a<br />

secondary ballis<strong>to</strong>spore), or by the formation of<br />

conidia. Repetitious germination is common in<br />

certain jelly fungi (Tremellales) (see Figs. 18.6b,<br />

21.7b and 21.13d). Germination by the formation<br />

of conidia is illustrated for Dacrymyces stillatus<br />

(Fig. 21.4a), Auricularia auricula-judae (Fig. 21.6c)<br />

and by yeast-like budding in Tremella frondosa (Fig.<br />

21.13c). During direct germination, germ tubes<br />

usually emerge through a special germ pore at<br />

the hilum or, as in Coprinus, through a pore at<br />

the opposite end of the spore.<br />

18.7.2 Monokaryotic and dikaryotic<br />

hyphae<br />

Because the nuclear divisions involved in basidiospore<br />

formation are meiotic, basidiospores<br />

are haploid. Since the post-meiotic nuclear<br />

divisions are mi<strong>to</strong>tic, if there are several nuclei<br />

in a single basidiospore these are usually<br />

genetically identical. They are said <strong>to</strong> be homokaryotic<br />

(Gr. homos ¼ equal, alike; karyon ¼ a nut,<br />

here meaning nucleus). At germination, repeated<br />

mi<strong>to</strong>tic nuclear divisions occur and the early<br />

germ tubes may consequently be multinucleate<br />

and coenocytic. Transverse septa are laid down<br />

behind the growing hyphal tip and eventually<br />

divide the hypha in<strong>to</strong> segments which contain<br />

only a single nucleus. The uninucleate segments<br />

and the hyphae which contain them are said <strong>to</strong><br />

be monokaryotic. The terms homokaryon and<br />

monokaryon or primary mycelium have also<br />

been applied <strong>to</strong> such haploid mycelia. As part of<br />

the sexual cycle, monokaryotic hyphae of genetically<br />

distinct mating type undergo plasmogamy<br />

(soma<strong>to</strong>gamy), i.e. they fuse <strong>to</strong>gether and initiate<br />

the formation of a mycelium made up of

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