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Introduction to Fungi, Third Edition

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170 ZYGOMYCOTA<br />

Fig 7.5 Sporangiophore of<br />

Phycomyces as a cylindrical lens.<br />

Upper portion: light ray L impinges<br />

from the left and is refracted at<br />

the first surface.The ratio of<br />

‘path length’of the light ray in<br />

the proximal part of the<br />

sporangiophore <strong>to</strong> the path length<br />

in the distal part is PM/DM.The<br />

maximum value of the angle b is<br />

about 20°.Lowerportion:<br />

sporangiophore in section <strong>to</strong> show<br />

the peripheral layer of cy<strong>to</strong>plasm<br />

surrounding the central vacuole.<br />

The values are estimates of the<br />

refractive index of cy<strong>to</strong>plasm and<br />

vacuolar sap. Diagram modified<br />

from Bergman et al. (1969).<br />

nature of the pho<strong>to</strong>recep<strong>to</strong>rs can be obtained by<br />

studying the action spectrum of the response<br />

over a range of light wavelengths. The pho<strong>to</strong>tropic<br />

curvature of Phycomyces sporangiophores<br />

has a similar action spectrum <strong>to</strong> the growth<br />

response of the vegetative mycelium, which is<br />

also stimulated by light. There are several clearly<br />

defined peaks at 485, 455, 385 and 280 nm, i.e.<br />

mostly in the blue part of the spectrum.<br />

Although b-carotene is present in large amounts<br />

in the growing zone of the sporangiophore, the<br />

pho<strong>to</strong>recep<strong>to</strong>r system is more likely <strong>to</strong> comprise<br />

a flavin-type molecule and a pterin-type protein<br />

(Flores et al., 1999; Galland & Tölle, 2003).<br />

Mutants with less than 0.1% of the wild-type b-<br />

carotene content remain fully pho<strong>to</strong>sensitive.<br />

However, b-carotene is involved in the other<br />

light-induced responses. The signalling chains<br />

involved in transduction of the light signal are<br />

only partially unravelled at present (Cerdá-<br />

Olmedo, 2001).<br />

Moss and Baker (2002) have described a<br />

technique for demonstrating the pho<strong>to</strong>tropic<br />

response of P. blakesleeanus in the labora<strong>to</strong>ry.<br />

7.2.3 Sporangiophore development in<br />

Phycomyces<br />

As the sporangiophore of Phycomyces develops it<br />

rotates. Castle (1942) followed the growth and<br />

rotation of the sporangiophore by attaching<br />

Lycopodium spores as markers and tracking the<br />

displacement of the markers. His findings are<br />

illustrated in Fig. 7.6. After a period of apical<br />

growth of the tubular sporangiophore (stage I),<br />

the sporangium appears as a terminal swelling<br />

and growth ceases. During this period (stage II),<br />

growth is limited <strong>to</strong> sporangial enlargement.<br />

In the next period (stage III) no further enlargement<br />

of the sporangium occurs, and elongation<br />

is also at a standstill. During stages IVA and IVB,<br />

elongation of the sporangiophore is resumed and<br />

growth is mainly localized in a zone somewhat<br />

below the sporangium. During stage I the tip of<br />

the sporangiophore rotates clockwise (as seen<br />

from above looking down) through a maximum<br />

angle of about 90°. There is no rotary movement<br />

during stages II and III. When sporangiophore<br />

elongation recommences in stage IVA, the direction<br />

of rotation is now anti-clockwise (as seen from

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