_aa._ =Hypothallus or Prothallus.= The first stage of growth in the
lichen thallus can be most easily traced in epilithic crustaceous
species, especially in those that inhabit a smooth rock surface. The
spore, on germination, produces a delicate branching septate mycelium
which radiates on all sides, as was so well observed and recorded by
Tulasne[317] in _Verrucaria muralis_ (Fig. 14). Zukal[318] has called
this first beginning the prothallus. In time the cell-walls of the
filaments become much thicker and though, in some species, they remain
colourless, in others they become dark-coloured, all except the extreme
tips, owing to the presence of lichen pigments—a provision, Zukal[319]
considers, to protect them against the ravages of insects, etc. The
prothallic filaments adhere closely to the substratum and the branching
becomes gradually more dendroid in form, though sometimes hyphae are
united into strands, or even form a kind of plectenchymatous tissue.
This purely hyphal stage may persist for long periods without much
change. In time there may be a fortuitous encounter with the algae (Fig.
38 A) which become the gonidia of the plant. Either these have been
already established on the substratum as free-growing organisms, or, as
accidentally conveyed, they alight on the prothallus. The contact between
alga and hypha excites both to active growth and to cell-division;
and the rapidly multiplying gonidia are as speedily surrounded by the
vigorously growing hyphal filaments.
[Illustration: Fig. 38 A. Hypothallus of _Rhizocarpon confervoides_ DC.,
from the extreme edge, with loose gonidia × 600.]
[Illustration: Fig. 38 B. Young thallus of _Rhizocarpon confervoides_
DC., with various centres of gonidial growth on the hypothallus × 30.]
Schwendener[320] has thus described the origin and further development
of prothallus and gonidia: on the dark-coloured proto- or prothallus,
he noted small nestling groups of green cells which he, at that time,
regarded as direct outgrowths from the lichen hyphae. These gonidial
cells, increasing by division, multiplied gradually and gathered into
a connected zone. He also observed that the hyphae in contact with the
gonidia became more thin-walled and produced many new branches. Some of
these newly formed branches grow upwards and form the cortex, others
grow downwards and build up the medulla or pith; the filaments at the
circumference continue to advance and may start new centres of gonidial
activity (Fig. 38 B). In many species, however, this prothallus or, as
it is usually termed at this stage, the hypothallus, becomes very soon
overgrown and obscured by the vigorous increase of the first formed
symbiotic tissue and can barely be seen as a white or dark line bordering
the thallus (Fig. 39). Schwendener[321] has stated that probably only
lichens that develop from the spore are distinguished by a protothallus,
and that those arising from soredia do not form these first creeping
filaments.
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