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LICHEN GONIDIA 21<br />

i. GONIDIA IN RELATION TO THE THALLUS<br />

A. HISTORICAL ACCOUNT OF LICHEN GONIDIA<br />

There have been few subjects of botanical investigation that have<br />

roused so much speculation and such prolonged controversy as the question<br />

of these constituents of the lichen plant. The green cells and the colourless<br />

filaments which together form the vegetative structure are so markedly<br />

dissimilar, that constant attempts have been made to explain the problem<br />

of their origin and function, and thereby to establish satisfactorily the<br />

relationship of lichens to other members of the Plant Kingdom.<br />

In gelatinous lichens, represented by Collema, of which several species<br />

are common in damp places and grow on trees or walls or on the ground,<br />

the chains of green cells interspersed through the thallus have long been<br />

recognized as comparable with the filaments of Nostoc, a blue-green<br />

gelatinous alga, conspicuous in wet weather in the same localities as those<br />

inhabited by Collema. So among early systematists, we find Ventenat 1<br />

classifying the few lichens with which he was acquainted under algae and<br />

hazarding the statement that a gelatinous lichen such as Collema was only<br />

a Nostoc changed in form. Some years later Cassini 2 in an account of Nostoc<br />

expressed a somewhat similar view, though with a difference: he suggested<br />

that Nostoc was but a monstrous form of Collema, his argument being that,<br />

as the latter bore the fruit, it was the normal and perfect condition of<br />

the plant. A few years later Agardh 3 claimed to have observed the meta-<br />

morphosis of Nostoc up to the fertile stage of a lichen, Collema limosum.<br />

But long before this date, Scopoli 4 had demonstrated a green colouring<br />

substance in non-gelatinous lichens by rubbing a crustaceous or leprose<br />

thallus between the fingers; and Persoon 5 made use of this green colour<br />

characteristic of lichen crusts to differentiate these plants from fungi.<br />

Sprengel 6 went a step further in exactly describing the green tissue as<br />

forming a definite layer below the upper cortex of foliaceous lichens.<br />

The first clear description and delimitation of the different elements<br />

composing the lichen thallus was, however, given by Wallroth 7 . He<br />

drew<br />

attention to the great similarity between the colourless filaments of the<br />

lichen and the hyphae of fungi. The green globose cells in the chlorophyllaceous<br />

lichens he interpreted as brood-cells or gonidia, regarding them as<br />

organs of reproduction collected into a "stratum gonimon." To the same<br />

author we owe the terms "homoiomerous" and "heteromerous," which he<br />

coined to describe the arrangement of these green cells in the tissue of the<br />

thallus. In the former case the gonidia are distributed equally through the<br />

structure; in the latter they are confined to a distinct zone.<br />

1 Ventenat 1794, p. 36.<br />

5 Persoon 1794, p. 17.<br />

2<br />

Cassini 1817, p. 395.<br />

3<br />

Agardh 1820.<br />

4<br />

Scopoli 1/60, p. 79.<br />

6<br />

Sprengel 1804, p. 325.<br />

7 Wallroth 1825, I.

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