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Introduction to Fungi, Third Edition

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188 ZYGOMYCOTA<br />

the sporangiophore of Pilobolus, as they do in<br />

many zygomycetes. Eventually, usually about<br />

midday, the sporangial vesicle explodes at a line<br />

of weakness just beneath the columella. Due <strong>to</strong><br />

the elasticity of the vesicle wall the liquid<br />

contents are squirted out, projecting the entire<br />

sporangium forward in the direction of the light.<br />

Pho<strong>to</strong>graphs of the jet show that it is at first<br />

cylindrical but eventually breaks up in<strong>to</strong> fine<br />

droplets (Fig. 7.22c; Page, 1964). In P. kleinii, the<br />

velocity of projection varies between wide limits<br />

of 4.7 27.5 m s 1 with a mean of 10.8 m s 1<br />

(Page & Kennedy, 1964). The sporangia can be<br />

projected vertically upwards for as much as<br />

2 m and horizontally for up <strong>to</strong> 2.5 m. On striking<br />

a grass blade or other herbage, the sporangium<br />

becomes attached in such a way that the<br />

mucilaginous ring adheres <strong>to</strong> it, with the black<br />

sporangium wall facing outwards. Buller (1934)<br />

has suggested that the projectile contains a<br />

drop of liquid attached <strong>to</strong> the sporangium<br />

(Fig. 7.22a). When the projectile strikes an<br />

object the liquid flows around the sporangium,<br />

but because the sporangium wall is hydrophobic<br />

and the base of the sporangium is surrounded<br />

by the wettable mucilaginous ring, the sporangium<br />

turns round in the liquid so that its wall<br />

faces outwards (Fig. 7.22b). The non-wettable<br />

nature of the sporangial wall may be related <strong>to</strong><br />

the presence on its surface of hollow, blunttipped<br />

spines and crystals as seen by electron<br />

microscopy (Bland & Charles, 1972). As the<br />

mucilage dries, the sporangium becomes<br />

cemented on<strong>to</strong> the surface which it struck. The<br />

spores of Pilobolus are released only after the<br />

sporangium has been ingested by an animal.<br />

They survive gut passage and are voided with<br />

the faeces. A film featuring the life cycle of<br />

Pilobolus has been made (Webster & Hard, 1999).<br />

An unexpected consequence of the attachment<br />

of Pilobolus sporangia <strong>to</strong> herbage is that the<br />

sporangia may act as vec<strong>to</strong>rs for parasitic<br />

nema<strong>to</strong>des such as Dictyocaulus spp., which multiply<br />

on dung and, when ingested, cause lungworm<br />

disease in sheep, cattle and some wild<br />

mammals.<br />

The physiology of Pilobolus shows a number<br />

of interesting features possibly related <strong>to</strong> its<br />

coprophilous habit. Spores germinate best above<br />

Fig 7.21 Pilobolus kleinii. Diagrammatic L.S. of sporangiophore<br />

showing the path of light rays falling parallel <strong>to</strong> the axis of the<br />

sporangiophore which are brought <strong>to</strong> a focus beneath the<br />

subsporangial vesicle.The sporangiophore illustrated is<br />

orientated symmetrically with respect <strong>to</strong> the incident light.<br />

Note the mucilaginous ring extruded through the sporangial<br />

wall at its base (after Buller,1934).<br />

pH 6.5, and can be induced <strong>to</strong> germinate by<br />

treatment with alkaline pancreatin. Germination<br />

can also be triggered by hexoses such as<br />

glucose and mannose. Mycelial growth occurs<br />

over a wide range of temperatures, with optimum<br />

temperatures at 25 35°C. Growth on<br />

synthetic media with asparagine and acetic acid

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