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Introduction to Fungi, Third Edition

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GENERAL ASPECTS OF LICHEN BIOLOGY<br />

451<br />

A pre-thallus can be formed by fusion of<br />

several genetically distinct isidia, soredia or<br />

pho<strong>to</strong>- and mycobionts, and likewise several<br />

pre-thalli can fuse in the process of thallus<br />

maturation. Thus, a mature thallus may contain<br />

a jigsaw of genetically heterogeneous myco- and<br />

pho<strong>to</strong>bionts (Fahselt, 1996). In other cases, e.g.<br />

the map lichen Rhizocarpon geographicum, the<br />

borders between incompatible thalli are demarcated<br />

by black barrage lines. In some lichens with<br />

a green alga as pho<strong>to</strong>biont, the situation is<br />

further complicated by the inclusion of a second<br />

(cyanobacterial) pho<strong>to</strong>biont. This is then usually<br />

confined <strong>to</strong> discrete regions termed cephalodia<br />

which often differ in their morphology from the<br />

thallus containing the green pho<strong>to</strong>biont (Fig.<br />

16.4).<br />

16.2.4 Lichenicolous fungi<br />

The capture of a compatible pho<strong>to</strong>biont partner<br />

by a germinating fungus spore can be problematic,<br />

especially if the pho<strong>to</strong>biont belongs <strong>to</strong> the<br />

genus Trebouxia which does not seem <strong>to</strong> be<br />

widespread as a free-living organism. One solution<br />

<strong>to</strong> the problem is the recovery of pho<strong>to</strong>bionts<br />

from the propagules of other lichen<br />

species. Certain pho<strong>to</strong>biont strains are favoured<br />

by many taxonomically unrelated mycobionts<br />

(Rikkinen et al., 2002). If these lichens grow<br />

in similar ecological situations, communities are<br />

formed which have a high mycobiont diversity<br />

but share the same or closely related pho<strong>to</strong>biont<br />

strains. Thus, the chances for a germinating<br />

spore <strong>to</strong> salvage compatible pho<strong>to</strong>bionts from<br />

soredia or isidia of other lichen species may be<br />

quite high (Rikkinen, 2003).<br />

Several mycobionts have taken the ultimate<br />

step of poaching their pho<strong>to</strong>biont from an<br />

existing lichen thallus in order <strong>to</strong> establish<br />

their own independent thallus (Ott et al., 1995).<br />

Such organisms are called lichenicolous lichens,<br />

and the phenomenon has been aptly named<br />

‘clep<strong>to</strong>biosis’ (Honegger, 1993). Numerous other<br />

fungi feed on the pho<strong>to</strong>synthetic products of a<br />

lichenized pho<strong>to</strong>biont without ever establishing<br />

an independent thallus, while yet others destructively<br />

parasitize the host lichen (Rambold<br />

& Triebel, 1992; Richardson, 1999; Lawrey &<br />

Diederich, 2003). Such fungi are collectively<br />

called lichenicolous fungi, and they are often<br />

taxonomically related <strong>to</strong> lichenized fungi.<br />

16.2.5 The nutritional basis of the<br />

lichen symbiosis<br />

Hill and Smith (1972) devised a simple and<br />

elegant method termed the ‘inhibition technique’<br />

which has permitted the identification of<br />

carbohydrates secreted by the pho<strong>to</strong>biont. Lichen<br />

thalli are exposed <strong>to</strong> radiolabelled CO 2 , and<br />

after a while an excess of a single unlabelled<br />

Fig16.4 Placopsisgelida, a foliose lichen.The<br />

brightly coloured main thallus containing the<br />

primary (algal) pho<strong>to</strong>biont has produced a<br />

dark central gall-like cephalodium in which<br />

the secondary (cyanobacterial) pho<strong>to</strong>biont is<br />

localized. From Bu«del and Scheidegger (1996),<br />

by permission of Cambridge University Press.<br />

Original image kindly provided by<br />

C. Scheidegger.

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