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Introduction to Fungi, Third Edition

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494 BASIDIOMYCOTA<br />

dikaryotic ballis<strong>to</strong>spore directly. A discharged<br />

primary ballis<strong>to</strong>spore may germinate by a germ<br />

tube or by repetition <strong>to</strong> form a secondary<br />

ballis<strong>to</strong>spore (Fig. 18.6b). Yeast-like growth can<br />

also occur, especially on rich media.<br />

Working with Itersonilia, several observations<br />

were made which provided clues <strong>to</strong> the mechanism<br />

of discharge. (1) Ballis<strong>to</strong>spores can be<br />

detached from their sterigmata with a micromanipula<strong>to</strong>r<br />

needle and spores so detached still<br />

develop Buller’s drop. This shows that the liquid<br />

in the drops does not originate from liquid<br />

transported through sterigmata. (2) During<br />

normal discharge, although the volume of<br />

Buller’s drop may attain 60% of that of the<br />

spore, there is no decrease in the dimensions of<br />

the ballis<strong>to</strong>spore, indicating that Buller’s drop<br />

does not come from within the spore. The same<br />

observation has been made on other basidiomycetes<br />

and has led <strong>to</strong> the suggestion that the<br />

liquid in Buller’s drop and also in the adaxial<br />

drop is formed by condensation of water vapour<br />

around a hygroscopic substance extruded from<br />

the hilar appendix and through the spore wall<br />

(Webster et al., 1984a,b, 1989). Washings from the<br />

spores of basidiomycetes belonging <strong>to</strong> several<br />

different taxonomic groups were analysed by<br />

gas liquid chroma<strong>to</strong>graphy, and all gave a positive<br />

result for the presence of manni<strong>to</strong>l. Glucose<br />

was also sometimes detected. The presence of<br />

manni<strong>to</strong>l and hexose in liquid drawn off by<br />

a micropipette from Buller’s drops in Itersonilia<br />

was confirmed by microscope fluorimetry, and<br />

measurements of the solute concentrations in<br />

Buller’s drop corresponded closely <strong>to</strong> the calculated<br />

concentrations which would be necessary<br />

<strong>to</strong> drive the uptake of water from a saturated<br />

atmosphere at the rates observed (Webster et al.,<br />

1995).<br />

The surface tension catapult mechanism<br />

postulates that, as Buller’s drop develops, the<br />

centre of mass of the spore plus drop moves<br />

<strong>to</strong>wards the hilar appendix (Fig. 18.8b). The<br />

coalescence of Buller’s drop with the adaxial<br />

drop causes a rapid redistribution of mass away<br />

from the hilar appendix, resulting in a momentum<br />

which carries the spore plus drop away from<br />

the sterigma (Fig. 18.8c; Webster et al., 1988).<br />

Pringle et al. (2005) have suggested that an even<br />

Fig18.6 Itersonilia perplexans. (a) Basidium (sporogenous cell)<br />

bearing a single ballis<strong>to</strong>spore. Note the clamp connection at<br />

the base of the basidium (arrow). (b) A primary ballis<strong>to</strong>spore<br />

has germinated by repetition <strong>to</strong> form a secondary<br />

ballis<strong>to</strong>spore.<br />

greater momentum may be generated by the<br />

fusion drop moving <strong>to</strong>wards the basidiospore<br />

apex and coming <strong>to</strong> an abrupt halt upon reaching<br />

it. For this mechanism <strong>to</strong> be effective, a rigid<br />

sterigma is required, and it is likely that the<br />

turgor pressure of the vacuolated basidium<br />

contributes <strong>to</strong> the required rigidity (Money, 1998).<br />

The requirement of high humidity for effective<br />

operation of the mechanism is a likely<br />

explanation for observations that basidiospore<br />

concentrations in the air peak at night (Kramer,<br />

1982). High humidity develops in the space<br />

between agaric gills or inside the hymenial<br />

tubes of <strong>to</strong>ads<strong>to</strong>ols and bracket fungi. The<br />

presence of free water would, of course, prevent<br />

operation of the surface tension catapult, and<br />

this may be the reason why agaric basidiocarps<br />

are often umbrella-shaped. The impossibility of<br />

the surface tension catapult mechanism operating<br />

under water also explains why, although<br />

some basidiomycetes grow vegetatively in fresh

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