If it be conceded that fully developed carpogonia with emergent
trichogynes, accompanied by spermogonia and spermatia, represent
fertilization, or the probability of fertilization, then the process may
be assumed to take place in a fairly large and widely distributed series
of lichens. Copulation between the spermatium and the trichogyne has
been seen by Stahl[650], Baur[651] and by F. Bachmann[652] in _Collema_.
In _Physcia pulverulenta_ Darbishire[653] could not prove copulation
in the earlier stages, but he found what he took to be the remains of
emptied spermatia adhering to the tips of old trichogynes. Changes in the
trichogyne following on presumed copulation have been demonstrated by
several workers in the Collemaceae, and open communication as a result of
fertilization between the cells of the ascogonium has been described in
two species. This coenocytic condition of the ascogonium (or archicarp),
considered by Darbishire and others as an evidence of fertilization, has
been demonstrated by Fitzpatrick[654] in the fungus _Rhizina undulata_.
The walls between the cells of the archicarp in that Ascomycete became
more or less open, so that the ascogenous hyphae growing from the central
cells were able easily to draw nutrition from the whole coenocyte, but
no process of fertilization in _Rhizina_ preceded the breaking down of
the septa and no fusion of nuclei was observed until the stage of ascus
formation.
The real distinction between fertile and vegetative hyphae lies,
according to Harper[655], in the relative size of the nuclei. F. Bachmann
speaks of one large nucleus in the spermatium of _Collema pulposum_ which
would indicate sexual function. There is however very little nuclear
history of lichens known at any stage until the beginning of ascus
formation, when fusion of two nuclei certainly take place as in fungi to
form the definitive nucleus of the ascus.
The whole matter may be summed up in Fünfstück’s[656] statement that:
“though research has proved as very probable that fertilization takes
place, it is an undoubted fact that no one has observed any such process.”
F. FINAL STAGES OF APOTHECIAL DEVELOPMENT
The emergence of the lichen apothecium from the thallus, and the form it
takes, are of special interest, as, though it is essentially fungal in
structure, it is subject to various modifications entailed by symbiosis.
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