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Introduction to Fungi, Third Edition

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450 LICHENIZED FUNGI (CHIEFLY HYMENOASCOMYCETES: LECANORALES)<br />

Fig16.2 Cladonia pyxidata. Soredia containing algal cells<br />

surrounded by fungal hyphae.<br />

Fig16.3 Interactions between pho<strong>to</strong>biontcells (Trebouxia) and mycobiont hyphae. (a,b) Early stages in the establishment of a lichen<br />

thallus in Cladonia cristatella. (a) Single algal cell at point of contact with the mycobiont. (b) Ensheathment of algal cell by the<br />

mycobiont. Both images redrawn and modified from SEM images of Ahmadjian and Jacobs (1981). (c) Formation of tubular<br />

intracellular haus<strong>to</strong>ria by the mycobiont of a simple crus<strong>to</strong>se lichen (e.g. Lecanora conizaeoides).The same pho<strong>to</strong>biont cell may be<br />

penetrated repeatedly. (d) Formation of an intraparietal haus<strong>to</strong>rium by the mycobiont of a stratified lichen. Exchange of nutrients<br />

occurs apoplastically; the walls of both mycobiont and pho<strong>to</strong>biont are surrounded by a common hydrophobin sheath produced by<br />

the former. (c,d) schematic drawings based on the results of Honegger (1986).<br />

If partners are compatible, a pre-thallus<br />

is formed, i.e. a crus<strong>to</strong>se non-stratified cluster<br />

of pho<strong>to</strong>biont cells ensheathed by hyphae of<br />

the mycobiont (Trembley et al., 2002). A prethallus<br />

is also formed by germinating isidia<br />

or soredia. Under suitable conditions, the prethallus<br />

enlarges and stratifies in<strong>to</strong> a mature<br />

thallus; the first sign of this step is the secretion<br />

of mucilage by hyphae at the periphery of the<br />

pre-thallus (Honegger, 1993).<br />

The fac<strong>to</strong>rs determining pho<strong>to</strong>biont<br />

mycobiont specificity and thus permitting the<br />

development of a pre-thallus and mature thallus<br />

are not yet known. In addition <strong>to</strong> genetic determinants,<br />

environmental fac<strong>to</strong>rs must also play<br />

a crucial role as indicated by the difficulties<br />

encountered when trying <strong>to</strong> grow pre-thalli<br />

in<strong>to</strong> fully differentiated thalli in the labora<strong>to</strong>ry<br />

(Ahmadjian, 1993). Recognition is probably a<br />

continuous and multi-step process mediated<br />

by surface molecules such as lectins, and facilitated<br />

by the embedding of both bionts in a<br />

gelatinous matrix of fungal origin (Ahmadjian,<br />

1993).

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