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Introduction to Fungi, Third Edition

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694 ANAMORPHIC FUNGI<br />

Bandoni (1974) has advanced the idea that<br />

tetraradiate spores may be adapted <strong>to</strong> movement<br />

in surface films of water.<br />

Although sigmoid spores are less efficiently<br />

trapped than tetraradiate spores, they, <strong>to</strong>o, can<br />

develop in different ways and in unrelated<br />

groups of fungi which have adopted an aquatic<br />

habit, suggesting that their shape has selective<br />

value. Observations on sigmoid spores moving<br />

with the current flow in flat capillary tubes show<br />

that as they approach a surface they tumble end<br />

over end and come <strong>to</strong> rest with one spore tip in<br />

contact with the surface. Immediately after<br />

arrest, the spore swings parallel <strong>to</strong> the current,<br />

thus minimizing the shear forces acting <strong>to</strong><br />

detach the spore (Webster & Davey, 1984). As in<br />

tetraradiate spores, the tips of the arms of<br />

sigmoid spores secrete mucilage, possibly in<br />

response <strong>to</strong> a thigmotropic stimulus associated<br />

with their tumbling movements. Mucilage is also<br />

present on the outside of the sigmoid spore of<br />

Mycocentrospora filiformis prior <strong>to</strong> it making<br />

contact with a surface (Au et al., 1996). Current<br />

flow forces the spore in<strong>to</strong> contact with the<br />

surface at a second point along its length.<br />

Germination occurs by the development of<br />

a germ tube from that end of the spore which<br />

is in contact with the surface. Thus, in contrast<br />

<strong>to</strong> the three-point contact associated with tetraradiate<br />

spore shape, sigmoid spores make two<br />

points of contact with surfaces <strong>to</strong> which they<br />

adhere.<br />

25.2.5 Spores in stream foam<br />

Foam is an effective trap for both tetraradiate<br />

and sigmoid spores (see Fig. 25.9). Spores<br />

suspended in water or caught in stream foam<br />

rarely germinate and many studies on the<br />

distribution and seasonal abundance of aquatic<br />

hyphomycetes using preserved foam samples<br />

have been made. In experiments in which air<br />

bubbles were passed through concentrated<br />

suspensions of conidia, the concentration of<br />

suspended spores fell very rapidly. Tetraradiate<br />

conidia were removed more readily than conidia<br />

of sigmoid or other shape (Iqbal & Webster,<br />

1973a). Comparisons of spores collected in foam<br />

or by Millipore filtration from the same stream<br />

indicates that the spore content of foam overrepresents<br />

the tetraradiate type of conidium<br />

in relation <strong>to</strong> other spore types known <strong>to</strong> be<br />

present. It cannot be assumed that all propagules<br />

found in stream foam originate from within<br />

the stream. Some come from fungi growing on<br />

the living leaves of riparian trees and the spores<br />

are brought in<strong>to</strong> the stream in raindrops or<br />

from rainwater draining down the tree trunks.<br />

Such fungi have been distinguished as terrestrial<br />

aquatic hyphomycetes (Ando & Tubaki,<br />

1984a,b).<br />

25.2.6 Adaptations of Ingoldian fungi <strong>to</strong><br />

the aquatic habitat<br />

Ingoldian fungi represent an ecological group of<br />

fungi sharing a common habitat, typically leaves<br />

and twigs in rapidly flowing streams. It is<br />

believed that they have been derived from<br />

terrestrial ances<strong>to</strong>rs and are able <strong>to</strong> colonize<br />

their habitat by virtue of a number of adaptations.<br />

These include effective spore attachment<br />

mechanisms associated with tetraradiate and<br />

sigmoid spore shape, rapid germination, and<br />

rapid growth and sporulation enhanced by<br />

turbulence and rapid water flow (Webster &<br />

Towfik, 1972; Sanders & Webster, 1980).<br />

Physiological adaptations include the possession<br />

of a range of enzymes enabling them <strong>to</strong> degrade<br />

their substrata (see below), and an ability <strong>to</strong><br />

grow at low temperatures, sometimes approaching<br />

0°C, so that they can continue growth<br />

and sporulation on submerged deciduous tree<br />

leaves after the autumn pulse of leaf fall<br />

(Koske & Duncan, 1974). Despite their typical<br />

environment, many aquatic hyphomycetes can<br />

survive for several weeks on dried leaves<br />

previously colonized in streams and brought<br />

out by flooding or by falling water levels (Sanders<br />

& Webster, 1978). It is possible that overland<br />

dispersal is achieved if colonized dried leaves<br />

are blown about by wind and re-deposited in<br />

streams.<br />

25.2.7 Ecophysiological studies<br />

There have been extensive studies on the ecology<br />

and physiology of Ingoldian hyphomycetes<br />

stimulated by the discovery that they play an

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