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LICHEN GONIDIA 27<br />

F. COMPOSITE NATURE OF THALLUS<br />

Schwendener 1 meanwhile was engaged on his study of lichen anatomy.<br />

Though at first he adhered to the then accepted view of the genetic connection<br />

between hyphae and gonidia, his continued examination of the<br />

vegetative development led him to publish a short paper 2 in which he<br />

announced his opinion that the various blue-green and green gonidia were<br />

really algae and that the complete lichen in all cases represented a fungus<br />

living parasitically on an alga: in Ephebe, for example, the alga was a form<br />

of Stigonema, in the Collemaceae it was a species of Nostoc. In those lichens<br />

enclosing bright green cells, the gonidia were identical with Cystococcus<br />

humicola, while in Graphideae the brightly coloured filamentous cells were<br />

those of Chroolepus (Trentepohlia). This statement he repeated in an<br />

appendix to the larger work on lichens 3 and again in the following year 4<br />

when he described more fully the different gonidial algae and the changes<br />

produced in their structure and habit by the action of the parasite: "though<br />

eventually the alga is destroyed," he writes, "it is at first excited to more<br />

vigorous growth by contact with the fungus, and in the course of generations<br />

may become changed beyond recognition both in size and form." In support<br />

of his theory of the composite constitution of the thallus, Schwendener<br />

pointed out the wide distribution and frequent occurrence in nature of the<br />

algae that become transformed to lichen gonidia. He claimed as further<br />

proof of the presence of two distinct organisms that, while the colourless<br />

filaments react in the same way as fungi on the application of iodine, the<br />

gonidia take the stain of algal membranes.<br />

G. SYNTHETIC CULTURES<br />

Schwendener's "dual hypothesis," as it was termed, excited great interest<br />

and no little controversy, the reasons for and against being debated with<br />

considerable heat. Rees 5 was the first who attempted to put the matte*. to<br />

the proof by making synthetic cultures. For this purpose he took spores<br />

from the apothecium of a Collema and sowed them on pure cultures of Nostoc,<br />

and as a result obtained the formation of a lichen thallus, though he did not<br />

succeed in producing any fructification. He observed further that the<br />

hyphal filaments from the germinating spore died off when no Nostoc was<br />

forthcoming.<br />

Bornet 6 followed with his record of successful cultures. He selected for<br />

experiment the spores of PJiyscia (Xanthoria) parietina and was able to<br />

show that hyphae produced from the germinating spore adhered to the free-<br />

1 Schwendener 1860, etc.<br />

4 Schwendener 1869.<br />

2 Schwendener 1867.<br />

5 Rees 1871.<br />

3 Schwendener 1868, p. 195.<br />

6 Bornet 1872.

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