_d._ PARASYMBIOSIS OF FUNGI. There occur on lichens, certain parasites
classed as fungi which at an early stage are more or less parasymbionts
of the host; as growth advances they may become parasitic and cause
serious damage, killing the tissues on which they have settled.
Zopf[950] found several instances of such parasymbiosis in his study
of fungal parasites, such as _Rhymbocarpus punctiformis_, a minute
Discomycete which inhabits the thallus of _Rhizocarpon geographicum_.
By means of staining reagents he was able to trace the course of the
parasitic hyphae, and found that they travelled towards the gonidia and
clasped them lichen-wise without damaging them, since these remained
green and capable of division. At no stage was any harm caused to the
host by the alien organism. Another instance he observed was that of
_Conida rubescens_ on the thallus of _Rhizocarpon epipolium_. By means
of fine sections through the apothecia of _Conida_ and the thallus of
the host, he proved the presence of numerous gonidia in the subhymenial
tissue, these being closely surrounded by the hyphae of the parasite,
and entirely undamaged: they retained their green colour, and in size
and form were unchanged. Zopf[951] at first described these parasites as
fungi though later[951] he allows that they may represent lower forms of
lichens.
Tobler[952] has added two more of these parasymbiotic species on the
border line between lichens and fungi, similar to those described
by Zopf. One of these, _Phacopsis vulpina_, belonging to the fungus
family Celidiaceae, is parasitic on _Letharia vulpina_. The fronds of
the host plant are considerably altered in form by its presence, being
more branched and curly. Where the parasite settles a swelling arises
filled with its hyphae, and the host gonidia almost disappear from the
immediate neighbourhood, only a few “nests” being found and these very
mucilaginous. These nests as well as single gonidia are surrounded by
_Phacopsis_ hyphae which have gradually displaced those of the _Letharia_
thallus. The gonidia are excited to division and increase in number on
contact with either lichen or fungus hyphae, but in the latter case the
increase is more abundant owing doubtless to a more powerful chemical
irritant in the fungus. As development advances, the _Phacopsis_ hyphae
multiply to the exclusion of both lichen hyphae and gonidia from the area
of invasion. Finally the host cortex is split, the fungus bursts through,
and the tissue beneath the parasite becomes brown and dead. _Phacopsis_
begins as a “parasymbiont,” then becomes parasitic, and is at last
saprophytic on the dead cells. The hyphae travel down into the medulla of
the host and also into the soredial outgrowths, and are dispersed along
with the host. The effect of _Verrucula_ on the host thallus may also be
cited[953].
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