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38<br />

CONSTITUENTS OF THE LICHEN THALLUS<br />

well as of colourless gonidia filled with the hyphal network. This description<br />

hardly tallies with the usual healthy appearance of the gonidial zone in the<br />

normal thallus, and it has been suggested that where the fungus filled the<br />

algal cell, it was as a saprophyte preying on dead material.<br />

The gradual perishing of algal cells in time by natural decay and their<br />

subsequent absorption by the fungus is undisputed. It is open to question<br />

whether the varying results recorded by these workers have any further<br />

significance.<br />

These observations of Elenkin and Danilov have been proved to be<br />

erroneous by Paulson and Somerville Hastings 1 . They examined the thalli<br />

of several lichens (Xanthoria parietina, Cladonia sp., etc.) collected in early<br />

spring when vegetative growth in these plants was found to be at its highest<br />

activity. They found an abundant increase of gonidia within the thallus,<br />

which they regarded as sporulation of the algae, and the most careful methods<br />

of staining failed to reveal any case of penetration of the gonidia by the<br />

hyphae.<br />

Nienburg 2 has published some recent observations on the association of<br />

the symbionts. In the wide cortex of a Pertusaria he found not only the<br />

densely compact hyphae, but also isolated gonidia. In front of these latter<br />

there was a small hollow cavity and, behind, parallel hyphae rich in contents.<br />

These gonidia had originated from the normal gonidial zone. They were<br />

moved upward by special hyphae called by Nienburg "push-hyphae." After<br />

their transportation, the algae at once divide and the products of division<br />

pass to a resting stage and become the centre of a new thalline growth. A<br />

somewhat similar process was noted towards the apex of Evernia furfur acea.<br />

Radial hyphae pushed up the cortex, leaving a hollow space over the gonidial<br />

zone. Into the space isolated algae were thrust by "push-hyphae." In this<br />

lichen he also observed the penetration of the algal cell by haustoria of the<br />

fungus. He considers that the alga reaps advantage but also suffers harm,<br />

and he proposes the term helotism to express the relationship.<br />

An instructive case of the true parasitism of a fungus on an alga has been<br />

described by Zukal 3 in the case of Endomyces scytonemata which he calls<br />

a "half-lichen." The mature fungus formed small swellings on the filaments<br />

of the Scytonema and, when examined, the hyphae were seen to have attacked<br />

the alga, penetrating the outer gelatinous sheath and then using up the<br />

contents of the green cells. It is only after the alga has been destroyed and<br />

absorbed, that asci are formed by the fungus. Zukal contrasts the development<br />

of this fungus with the symbiotic growth of the two constituents in<br />

EpJiebe where both grow together for an indefinite time.<br />

Mere associated growth however even between a fungus and an alga<br />

does not constitute a lichen. An instance of such growth is described by<br />

1 Paulson and Hastings 1920.<br />

2<br />

Nienburg 1917.<br />

3 Zukaj l89I>

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