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Introduction to Fungi, Third Edition

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ZYGOMYCETES: MUCORALES<br />

175<br />

but is converted <strong>to</strong> the active hormone trisporic<br />

acid after diffusion in<strong>to</strong> the mycelium of the<br />

complementary strain (Gooday, 1994). Trisporic<br />

acid stimulates further synthesis of b-carotene<br />

and of the two prohormones, leading <strong>to</strong> amplification<br />

of its own synthesis by a ‘cascade’<br />

mechanism. The 15 20-fold enhanced synthesis<br />

of b-carotene upon mating of two compatible<br />

strains of B. trispora holds potential for commercial<br />

production of this substance (Lampila et al.,<br />

1985; Sandmann & Misawa, 2002).<br />

The role of the trisporic acid in inducing<br />

b-carotene synthesis and zygophore formation is<br />

widespread in the Mucorales, having been<br />

characterized in Blakeslea, Phycomyces, Mucor and<br />

even Mortierella (Schimek et al., 2003). It is also<br />

known that trisporic acid is involved in the<br />

sexual response of some homothallic Mucorales<br />

such as Zygorhynchus moelleri, Mucor genevensis and<br />

Syzygites megalocarpus (Lampila et al., 1985). The<br />

common nature of the hormones of homothallic<br />

and heterothallic species could also be inferred<br />

from earlier observations of attempted matings<br />

between such forms, either at the interspecific or<br />

intergeneric level.<br />

Zygophores show directional growth <strong>to</strong>wards<br />

each other in response <strong>to</strong> volatile hormones.<br />

Gooday (1994) has suggested that these may be<br />

the mating type-specific prohormones, methyl-<br />

4-dihydrosporate of the (þ) strain and trisporol of<br />

the ( ) strain.<br />

Thigmotropic reactions<br />

When compatible zygophores make contact, they<br />

become firmly attached <strong>to</strong> each other and<br />

develop in<strong>to</strong> progametangia. In Mucor mucedo<br />

there is evidence that the cell wall chemistry of<br />

the zygophores is distinct from that of the<br />

vegetative mycelium and that the (þ) and ( )<br />

zygophores are bound <strong>to</strong>gether by lectins, i.e.<br />

glycoproteins exhibiting specific binding for<br />

polysaccharides (Jones & Gooday, 1977). In<br />

Fig 7.9 Collaborative biosynthesis of trisporic acid<br />

by cross-feeding of intermediates between (þ)and<br />

( ) mating types of Blakeslea trispora. b-Carotene<br />

(I) is metabolizedby both (þ)and( ) mating-types<br />

via retinal (II) <strong>to</strong> 4-dihydrotrisporol (III).This is<br />

metabolized by (þ)strains<strong>to</strong><br />

4-dehydrosporic acid and its methyl ester (IV)and<br />

by ( ) strains <strong>to</strong> trisporol (V).These are converted<br />

<strong>to</strong> trisporic acid (VI) only after diffusing <strong>to</strong> the ( )<br />

and (þ) strains, respectively. Redrawn from<br />

Gooday (1994), with kind permission of Springer<br />

Science and Business Media.

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